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N.C. DOCUMENTS
CLEAPIMGHOUSe
AUG 2 6 1998
r»if. immt 8F NORTH CAROLINA
MUltH
number 25 july 1998
EDITORIAL STAFF
Richard A. Lancia, Editor
Suzanne A. Fischer, Assistant Editor
Stephen D. Busack, Managing Editor
Brimleyana, the Zoological Journal of the North Carolina State Muse-um
of Natural Sciences, appears twice yearly in consecutively numbered issues.
Subject matter focuses on systematics, evolution, zoogeography, ecology, behav-ior,
and paleozoology in the southeastern United States. Papers stress the results
of original empirical field studies, but synthesizing reviews and papers of signif-icant
historical interest to southeastern zoology are also included. Brief com-munications
are accepted.
All manuscripts are peer reviewed by specialists in the Southeast and
elsewhere; final acceptability is determined by the Editor. Address manuscripts
and related correspondence to Editor, Brimleyana, North Carolina State Museum
of Natural Sciences, P.O. Box 29555, Raleigh, NC 27626. Information for con-tributors
appears in the inside back cover.
Address correspondence pertaining to subscriptions, back issues, and
exchanges to Brimleyana Secretary, North Carolina State Museum of Natural
Sciences, P.O. Box 29555, Raleigh, NC 27626.
In citations, please use the full name - Brimleyana.
North Carolina State Museum of Natural Sciences
Betsy Bennett, Director
North Carolina Department of Environment,
Health, and Natural Resources
James B. Hunt, Jr., Governor
Jonathon B. Howes, Secretary
CODN BRIMD 7
ISSN 0193-4406
IN MEMORIAM
Dr. Joshua Laerm, Professor at the University of Georgia and Curator of
Zoological Collections at the University of Georgia Museum of Natural History, died
28 September 1997. He was bom and raised in Pennsylvania and received an under-graduate
degree from Pennsylvania State University and graduate degrees from the
University of Illinois. He joined the faculty at the University of Georgia in 1976.
Dr. Laerm published numerous works in systematics, mammalogy, and nat-ural
history. He was particularly interested in rare and threatened or endangered mam-mals
and contributed significantly to understanding their natural history and distribu-tion
in the Southern Appalachian Mountains. His enthusiasm for science, his prolific
contributions, and his eagerness to help colleagues will be deeply missed.
Turtles (Reptilia: Testudines) Of The Ardis Local Fauna Late
Pleistocene (Rancholabrean) Of South Carolina
Curtis C Bentley and James L. Knight
South Carolina State Museum, 301 Gervais Street,
P.O. Box 100107
Columbia, South Carolina 29202-3107
ABSTRACT- The Ardis local fauna (late Pleistocene) was collected from
a group of interconnecting sediment-filled solution cavities, located in
the Giant Cement Quarry near Harleyville, Dorchester County, South
Carolina. Fossil material from the lowermost levels and the extreme
upper layer of the deposit have been radiocarbon dated at 18,940 ± 760
and 18,530 ± 725 y.b.p., respectively. These dates are considered con-temporaneous
within present resolution. Approximately ninety verte-brate
taxa were collected from the site. Fourteen were species of turtles,
including eight not previously reported from the Pleistocene of South
Carolina. Among these is the southeasternmost occurrence of Emy-doidea
blandingii. This record, in conjunction with other vertebrate fos-sils
from the site, suggests a north-south dispersal route of species along
the Atlantic Coastal Plain during interglacial-glacial transitions. Geo-graphically
isolated eastern and western populations of Emydoidea
blandingii may have existed during the maximum advance of the Lau-rentide
ice sheet. Unusually complete fossils of large box turtles recov-ered
from the site corroborate the previously suggested synonymy of the
extinct Terrapene Carolina putnami with T c. major. The fossil turtle
community of the Ardis local fauna has no modern analogue. Like the
Ardis mammals, it comprises a "disharmonious" fauna which suggests
that, during the height of the Wisconsinan glaciation, the region experi-enced
a more equable climate than that of today .
The Ardis local fauna has yielded approximately 90 species of late
Pleistocene fossil vertebrates from the Coastal Plain of South Carolina, includ-ing
a substantial mammalian (Bentley et al. 1994), avian, reptilian, amphibian,
and fish faunas currently under study. We present data on the Ardis turtle col-lection
(Appendix I), the largest Rancholabrean fauna reported from the state.
Dobie and Jackson (1979) and Roth and Laerm (1980) reported fossils of late
Pleistocene age from Edisto Island, the only other Pleistocene fossil turtle fauna
from South Carolina described to date. Among the Edisto Island fauna were ten
Curtis C. Bentley and James L. Knight
taxa of turtles, including Gopherus sp. and Malaclemys terrapin which were not
recovered from the Ardis local fauna and possibly three species of Pseudemys, P.
floridana and/or P. concinna, and P nelsoni.
The Ardis local fauna was discovered in a large open-pit mine, operat-ed
by the Giant Cement Company, located 5 km NNE of Harleyville, Dorchester
County, South Carolina (33° 14'N, 80° 26'W). Quarry operations exposed San-tee
Limestone (middle Eocene) and the clay-rich Harleyville Formation (late
Eocene) which underlie Plio-Pleistocene surficial deposits (Ward et al. 1979,
Harris and Zullo 1991). Locally, groundwater differentially dissolved the Santee
Limestone in its upper portions, so that many solution cavities contacted and
penetrated the overlying Harleyville Formation and thereby opened several of
the cavities to the Pleistocene surface (Bentley et al, 1994). The radiocarbon
dates of the Ardis material place the time of deposition at or near the height of
the Wisconsinan glaciation (Bowen 1988, Tushingham and Peltier 1993). Fur-ther
discussion on the geology, dating methods of the Ardis fossil material, pre-vious
fossil collections from the quarry, fossil collection procedures, and a local-ity
map are available in prior publications (Bentley and Knight 1993, Bentley et
al. 1994).
TAPHONOMY
At least part of the fossil assemblage collected from inside the solution
cavities at the Ardis site appears to represent an obrution deposit, the very rapid
burial of intact organisms (Brett 1990) in which many of the specimens exhibit
incipient decay. Surface openings leading to the cavities varied from a gentle
downward slope to a vertical shaft, generally allowing the Pleistocene fauna ease
of ingress and egress. This permitted animals to enter the cavities in three dif-ferent
ways: (1) "walk-in" taxa, which may have used the site for
estivation/hibernation or as denning sites and hunting grounds, for example
muskrats, mink, and woodrats (Bentley et al. 1994); (2) "wash-in" taxa from the
surface, either alive or dead, which applies most readily to large animals known
only from isolated remains e.g., Mammut sp., Bison sp., Equus sp. (Bentley et al.
1994), that would have been unable to enter the cavities during life; and (3) "fall-in"
taxa which fell into exposed verticle shafts, fossil accumulations resulting
from this type of natural trap are well documented (e.g., Webb 1974).
Because of the interconnecting "tunnel-like" nature of the cavities, a
single episodic event could produce differing water velocities within the cavities
and different rates of deposition. Seasonal flooding, depending on the intensity,
may have simultaneously smothered living animals within the cavities and
buried or reworked those that had died just prior to, or in a preceding, deposi-tional
event. Consequently, specimens incompletely or shallowly buried during
an event with a low sedimentation rate (low energy) could be completely or par-tially
exhumed and reburied by a succeeding event. This resulted in the preser-
Turtles
vation of specimens in various orientations to the bedding planes (Fig. 1), vari-ous
degrees of disarticulation, and the occasional mixing of individual elements.
An articulated Emydoidea blandingii specimen preserved in its life position with
axial skeletal elements inside, indicates little or no decay prior to its final burial
(Fig. 2). This preservation suggests the turtle was buried quickly in a high ener-gy,
high sedimentation environment (Brett and Speyer 1990), resulting in burial
deep enough to avoid reworking during subsequent episodic events. Several
articulated turtles were collected with limbs and skulls preserved within the
shells in various orientations to the bedding plane. A high energy hydrological
environment before or shortly after death would likely explain the various orien-tations
observed in well preserved specimens. Retention and preservation of
limb elements, cervical and caudal vertebrae, and skulls inside the shell may
reflect a withdrawal by the turtles in response to a catastrophic event.
Fig. 1 Emydoidea blandingii, only the carapace (.547) was preserved, ventral
side up (side view), among clay clasts from the surrounding Harleyville Forma-tion.
This illustrates the hydrodynamic effect upon some specimens prior to final
burial.
Curtis C. Bentley and James L. Knight
Fig. 2 Complete E. blandingii (.546) in situ, with axial skeleton preserved inside
the shell, indicating a "withdrawal response" and final burial prior to any signif-icant
decay.
MATERIALS AND METHODS
Morphological terminology used in this paper is taken from Carr
(1952), Ernst and Barbour (1989), Holman (1967, 1977, 1985), Holman and
Grady (1987), and Preston (1979). Taxonomy follows Conant and Collins
(1991).
Morphological comparisons of Recent skeletons to fossil material were
made against available specimens in the Florida Museum of Natural History and
the South Carolina State Museum collections. Additionally, specimens from The
University of Michigan Museum of Zoology of Emydoidea blandingii, UMMZ
155047-155054, Clemmys guttata, UMMZ 51235, 51236, 51240-51242,
159219, 155001, 155002, and Clemmys muhlenbergii, UMMZ 77140 and,
130840 were studied.
Most of the specimens in the South Carolina State Museum collections
are deposited under the base number of S.C. 94.10. and for brevity, are refer-enced
in the text only by the digits following this base number. Specimens
accessioned separately are designated by the institutional prefix of SCSM. Fos-sil
specimens deposited in the National Museum of Natural History and the
Florida Museum of Natural History are designated by USNM and UF, respec-tively.
Turtles
SYSTEMATIC PALEONTOLOGY
Testudines
Kinosternidae
Material: 1 right xiphiplastron (.25); 16 peripherals (.26-.44); 2 humeri (.45-. 46);
2 partial jaw rami (.755-. 756).
Remarks: These fossil elements could only be identified with confidence to fam-ily.
Kinosternon subrubrum - Eastern Mud turtle (Lacepede, 1788)
Material: 3 nuchals (.11 -.13); 2 right, 3 left hyoplastra (.6-. 10); 5 left hypoplas-ia
(.1-.5).
Characters used for identification: The hyoplastron of Kinosternon subrubrum
can be separated from other North American Kinosternon and Sternotherus
because the axillary notch is narrower, and from Kinosternon baurii because the
axillary notch is wider and shallower (Holman 1985). K. subrubrum hyo- and
hypoplasia differ from Sternotherus odoratus in that the elements are shorter lat-erally
than medially in S. odoratus (Preston 1979). Characters used to identify
nuchal material are discussed by Holman (1975). In addition, nuchals of K. sub-rubrum
can be distinguished from nuchals of S. odoratus because the anterior lip
of the nuchal, viewed anteriorly, is nearly straight in K. subrubrum. Nuchals of
S. odoratus, viewed anteriorly, have a decided arc.
Remarks: The eastern mud turtle inhabits a variety of shallow slow to non-mov-ing
bodies of water with a soft substrate, such as swamps, ponds, marshes, wet
meadows, and lagoons (Ernst and Barbour 1989). Kinosternon subrubrum today
ranges from southern Massachusetts and Pennsylvania along the Atlantic coast,
to the tip of Florida and west into Texas and Oklahoma (Conant and Collins
1991). K. subrubrum is common in the area of the Ardis site today and may be
sympatric with K. baurii (Lamb and Lovich 1990).
This is the first report of this species from the fossil record of South
Carolina. Dobie and Jackson (1979) and Roth and Laerm (1980) both reported
the same single pygal bone from Edisto Island as "Kinosternon sp."
Sternotherus odoratus - Common Musk turtle (Latreille, in Sonnini and
Latreille, 1802)
8 Curtis C. Bentley and James L. Knight
Material: 3 nuchals (.22-.24); 2 right hyoplastra (.14-. 15); 1 right, 5 left
hypoplasia (.16-. 21).
Characters used for identification: Identification is based on characters provid-ed
in discussion for Kinosternon subrubrum. The hyo-hypoplastron of S. minor
can be separated from the same elements in S. odoratus because the area that
forms the bridge between the plastron and the carapace is dorsally compressed
or flattened in S. minor, and not raised as in S. odoratus. The nuchals compare
most favorably to S. odoratus, following the characters used above, and addi-tionally
exhibit a strong dorsal medial keel that is generally lacking in K. sub-rubrum.
Remarks: The common musk turtle inhabits areas very similar to that of K. sub-rubrum,
preferring slow to non-moving bodies of water with a soft bottom. It
has also been collected from fast moving, gravel bottomed, streams (Ernst and
Barbour 1989). S. odoratus occurs from southern Maine and Canada southward
through Florida and as far west as Kansas and central Texas (Conant and Collins
1991), and occurs in the area of the Ardis site today.
Chelydridae
Chelydra serpentina - Snapping turtle (Linnaeus, 1758)
Material: 2 right parietals (.50, .53); 1 left postorbital (.51); 1 right prefrontal
(.52); 1 left quadratojugal (.54); 1 partial left mandible (.55); 2 right mandibles
C56-.57); 6 cervical vertebrae (.102-. 107); 1 humerus (.63); 2 radii (.69-.70); 1
right scapulo-acromial process (.64); 1 right partial acromial process (.65); 5
femora (.58-.62); 3 ilia (.66-.68); 1 caudal vertebra (.108); 1 nuchal (.95); 1 right
1st peripheral (.71); 16 unassigned peripherals (.72-.83)(2 USNM)(2 UF); 1 left
1st costal (.84); 24 partial costals (.85-.92)(8 USNM)(8 UF); 2 associated costals
C93-.94); 2 neurals (.96-.97); 2 epiplastra (.100-. 101); 2 hypoplasia (.98-.99).
Characters used for identification: Chelydra shell material is very distinctive and
easily separated from other turtles including Macroclemys. Preston (1979) pro-vides
characters that allow the identification of fragmentary material. All listed
fossil elements compare favorably to Recent skeletal materials.
Axial and appendicular skeleton - The large size and diagnostic orna-mentation
of the Chelydra skull roof elements distinguish them from all other
turtles. Macroclemys lacks the rugose cranial ornamentation of Chelydra. A
pectoral girdle was assigned to this species based on the 90° angle between the
scapula and acromial process and on the heavily striated distal ends (Holman
1966). Femora and humeri could not be separated from Recent material of C.
serpentina, and are more robust than other genera of fresh water turtles (Holman
1964) except Macroclemys, which is generally considerably larger.
Turtles
Remarks: Snapping turtles are generally found in freshwater to brackish water
habitats with soft muddy substrate (Ernst and Barbour 1989) from eastern Cana-da
through the United States east of the Rockies south through Mexico and into
Ecuador. C. serpentina occurs today in the Ardis area.
Dobie and Jackson (1979), first reported fossil material of C. serpenti-na
from Edisto Island with additional material reported by Roth and Laerm
(1980).
Macroclemys temminckii - Alligator snapping turtle (Gray, 1855)
Material: 1 partial right parietal (.109).
Characters used for identification: The fossil parietal is identical to Recent spec-imens
of this turtle, differing from C. serpentina in that the dorsal surface is
smooth, i.e. without any of the prominent ornamentation consistently found in C.
serpentina (Fig. 3). The parietal of M. temminckii is generally more robust and
is longer with respect to width than specimens of C. serpentina of comparable
sizes. This was the only fossil element of this species collected from the site.
Because this represents a significant range extension, assignment to this species
was made only after exhaustive comparisons to the fossil and Recent collections
at the Florida Museum of Natural History and the South Carolina State Museum
negated all other possibilities.
Remarks: This is the largest freshwater turtle in North America and possibly the
heaviest in the world (Ernst and Barbour 1989). The alligator snapping turtle
often can be found in the deep waters of lakes, ponds, rivers and bayous that con-tain
abundant aquatic vegetation and muddy bottoms. This turtle is highly aquat-ic
and ranges westward from northern Florida into Texas along the Gulf Coast
and thence northward up the Mississippi Valley into Illinois, Iowa and Kansas
(Ernst and Barbour 1989).
This is the first fossil or Recent evidence of Macroclemys from South
Carolina and the Atlantic Coastal Plain.
Emydidae
Emydinae
Chrysemys picta - Painted turtle (Schneider, 1783)
Material: An individual specimen consisting of a complete carapace (missing the
3rd right marginal) and plastron (.110); 7 cervical vertebrae (.110.1 -.110.7); 1
ulna (.110.16); 1 radius (.110.17); 4 phalanges (.110.18-.110.21); 1 ungual
(.110.22); 2 partial scapulo-acromial processes (.11 0.11 -.11 0.1 2); 2 coracoids
10 Curtis C. Bentley and James L. Knight
Fig. 3 Right parietals of Macroclemys temminckii and Chelydra serpentina. A)
Recent M. temminckii. B) Ardis fossil. C) Recent C. serpentina.
3 cm
(.1 10.13-.1 10.14); 2 dorsal vertebrae (.1 10.9-. 1 10.10); complete pelvic girdle
(.110.15); 1 caudal vertebra (.110.8). An individual specimen consisting of a
nearly complete carapace and plastron (.111). An individual specimen consist-ing
of a complete carapace (missing left 8-9th peripherals and 6th neural) and
plastron (.112); partial skull and mandible (.112.1); partial hyoid process
(.11 2.2); 7 cervical vertebrae (.112.20-. 11 2.26); 2 humeri (.11 2.5-. 11 2.6); 2 ulnae
(.112.7-.112.8); 2 radii (.112.9-. 112. 10); 2 scapulo-acromial processes (.112.15-
.112.16); 2 coracoids (.112.17-.112.18); 2 femora (.112.3-.112.4); 2 tibiae
(.112.11-.112.12); 2 fibulae (.112. 13-. 112. 14); 7 metapodial elements (.112.42-
.112.48); 31 phalanges (.112.49-. 112-80); 13 unguals (.112.81-.112.91); 2 sacral
ribs (.112.92-. 112.93); partial pelvic girdle (.112.19); 3 dorsal vertebrae (.112.27-
.112.29); 12 caudal vertebrae (.112.30-. 11 2.41). An individual specimen con-sisting
of a nearly complete plastron and attached 4-7th right peripherals (.113);
skull (including inner ear ossicles) and mandible (.113.1); partial hyoid appara-tus
(.113.2); 4 cervical vertebrae (.113-.9-.113.12); 2 humeri (.113.3-.113.4); 1
scapulo-acromial process (.113.7); 2 partial femora (.11 3.5-. 11 3.6); 1 tibia
(.113.8); 2 metapodial elements (. 1 1 3. 15-. 1 13. 16); 15 phalanges (.113.17-
.113.31); 5 unguals (1 13.32-. 1 13.36); 2 caudal vertebrae (. 1 13. 13-. 1 13. 14).
Specimen with partial carapace (.114); 3 cervical vertebrae (.114.3-. 114.6); 1
Turtles 11
humerus (.114.1); 1 radius (.114.2); 1 phalange (.114.7). 2 mixed specimens
both with partial fragmented carapaces and plastra (.115); 3 cervical vertebrae
(.115.5-. 115.6); 2 humeri (.115.1 -.115.2); 2 scapulo-acromial processes (.115.3-
.113.4); 2 coracoids (. 1 15.8-. 1 15.9); 2 tibiae (.115.10-.115.il); 1 ilium (.115.12);
1 phalange (.115.13). Four individual specimens with fragmented carapace and
plastron (.116-. 119). Two mixed specimens with badly fragmented carapaces
and plastra (.120).
Isolated elements: 16 nuchals (.123-.132)(3 USNM)(3 UF); (SCSM
91.170.1) partial plastron; 10 right and 2 left epiplastra (.133-. 144); 2 entoplas-tra
(.169-.170); 10 left and 3 right hyoplastra (.145-.151)(3 USNM)(3 UF); 1
right hypoplastron and xiphiplastron (.122); 5 left and 7 right hypoplasia (.152-
.157)(3 USNM)(3 UF); 6 left and 9 right xiphiplastra (.158-.168)(2 USNM)(2
UF); 8 right and 6 left 2nd costals (.227-.240); 3 right and 1 left 3rd costals (.241-
.244); 6 right and 3 left 4th costals (.245-.253); 7 right and 8 left 5th costals
C254-.268); 4 right and 3 left 6th costal (.269-.275); 4 right and 1 left 7th costal
(.276-.280); 56 peripherals (.171-.226); 1 mandible (.761); 2 right and 1 left
mandibular rami (.758-.760); 10 humeri (.289-.298); 8 femora (.281-.288).
Characters used for identification: Identification of complete shells was based
on nuchal characters (Bentley and Knight 1993), and the alignment of the verte-bral
and pleural sulci (Ernst and Barbour 1989) (Fig. 4).
Hyoplastron - The humeral sulcus does not cross dorsally over the plas-tral
scute overlap area, as in Clemmys. Terrapene and Emydoidea have hinged
plastra. Elements of comparable size can be separated from Deirochelys reticu-laria
as the dorsal scute overlap area in C. picta is wider and more sharply curved
Fig. 4 Fossil carapace of Chrysemys picta (.110) from the Ardis local fauna.
1 2 Curtis C. Bentley and James L. Knight
distally, and the articulating surface of the epiplastron and hyoplastron between
the entoplastron and the outside edge is wider in C. picta. Trachemys and
Pseudemys are larger and more robust than C picta as adults. Young specimens
of Trachemys and Pseudemys can be separated by a less pronounced or inflated
scute overlap area and signs of incomplete ossification.
Hypoplastron - The inguinal sulcus runs diagonally to the peripherals
and into the inguinal notch in Chrysemys but is parallel to the peripherals in
Clemmys. This element can be distinguished from Deirochelys as it is less elon-gate
with respect to width in C picta, and the scute overlap area is wider. It can
also be separated from Terrapene Carolina and Emydoidea by the lack of a hinge.
Adult Trachemys and Pseudemys differ in being larger and more robust than
adults of C. picta. Young specimens of Trachemys, comparable in size to adult
C. picta and completely ossified, can be separated from C. picta by a larger
bridge with respect to the hypoplastron proper and a greatly reduced or absent
epidermal attachment scar.
Entoplastron - The humero-pectoral sulcus does not cross the entoplas-tron
as in eastern species of Clemmys and in Terrapene Carolina. It can be tenta-tively
separated from Pseudemys, Trachemys, and Emydoidea by overall size, as
specimens of the preceding genera tend to exhibit incomplete ossification when
of comparable size to adult C. picta. However, size alone is not a reliable char-acter
for this element. We were unable to separate this element from that of
Deirochelys, so entoplastra are only tentatively assigned to C. picta.
Epiplastron - This element differs from other species (except Trache-mys)
in that the anterior edge exhibits a degree of serration. This element often
is serrated in specimens of Trachemys, but the size of these specimens allows
easy separation from C. picta. Young specimens of Trachemys generally lack
this serration and have a poorly developed scute overlap area in comparison to
C. picta of comparable size.
Xiphiplastron - This element can be separated from Clemmys, Ter-rapene,
and Emydoidea by the scute overlap area, which in those genera is more
pronounced than in C. picta. Further, Clemmys muhlenbergii has a posterior
edge tapered to a point, while in Clemmys insculpta the element is longer with
respect to width, with a pronounced notch where the anal sulcus wraps over the
edge, a condition minimal or lacking in C. picta. In C picta the scute overlap of
this element is wider and more pronounced on the posterior edge than on any
examined specimens of Deirochelys. This element in Trachemys is generally
more robust in adult specimens than in C. picta, and in young specimens of com-parable
sizes the scute overlap area is much less pronounced than that of C picta.
2nd Costal - This element differs from C guttata in being approxi-mately
30% wider with respect to length, and the junction between the 2nd ver-tebral
sulcus and the 1st and 2nd pleural sulcus is located generally more distal-ly
than in C. guttata. This element differs from C. muhlenbergii by having a
Turtles 1
3
greater curvature, and the above mentioned junction point forms a distinct "T"
shape in C. muhlenbergii and dips into a general "U" or "V" shape in C. picta.
The 2nd costal of C. picta differs from C. insculpta by being completely smooth
and lacking any "tortoise-like" bulges, by being more distally flared, and by hav-ing
greater curvature of the element. It can be separated from Trachemys and
Pseudemys by a lack of sculpting and smaller size, and from Deirochelys by lack
of sculpting, a more proximal rib attachment (Jackson 1978), and the proximal
tapering of the element.
3rd Costal - Differs from Clemmys, Terrapene, Trachemys, Pseudemys,
Emydoidea, and Deirochelys in that it lacks the vertebral sulcus between the 2nd
and 3rd vertebral scutes. Additionally, in Deirochelys the rib attachment is sig-nificantly
more distal than in C picta. This 3rd element can be separated from
the 5th costal in C. picta in that it lacks the posteriorly directed curvature of the
5th costal.
4th Costal - Can be separated from all other emydid turtles in that it
exhibits alignment of the sulci between the 2nd and 3rd vertebral and pleural
scutes. This sulcal alignment occurs only in the subspecies C picta picta
(Conant and Collins 1991).
5th Costal - Can be separated from Clemmys, Trachemys, Deirochelys,
Pseudemys, Emydoidea, and Terrapene by characters given for the 3rd costal.
6th Costal - Differs from Clemmys, Trachemys, Deirochelys, Pseude-mys,
Emydoidea, and Terrapene in that it lacks the sulcus of the 3rd and 4th ver-tebral
scutes.
Peripherals, femora, humeri, and mandibular rami - These elements are
tentatively assigned to this species as they compare most favorably to Recent and
fossil material of C. picta.
Remarks: The painted turtle has a wide distribution, occurring from southern
Canada south into Mexico and across the entire continental United States (Ernst
and Barbour 1989). Lakes, ponds, and streams are typical habitats of the paint-ed
turtle. Slow to non-moving, shallow aquatic environments with soft bottoms
are favored.
The completeness of many of the painted turtles recovered from the
Ardis site are strong indicators of an obrution deposit. Chrysemys picta occurs
in the Piedmont and mountains of South Carolina today but does not inhabit the
Ardis site or any other part of the Coastal Plain. This is the first fossil record
from South Carolina.
Clemmys guttata - Spotted turtle (Schneider, 1792)
Material: An individual specimen consisting of a nearly complete shell (SCSM
93.90.1) missing only the right hypoplastron and xiphiplastron, figured in Bent-
1 4 Curtis C. Bentley and James L. Knight
ley and Knight (1993), and the following associated cranial and postcranial ele-ments;
articulated skull fragment (prefrontal, frontal, postorbital, parietal, and
supraoccipital), right maxilla, both quadrates, both opisthotics, basisphenoid, and
articulated lower jaw, 1st cervical vertebra, 2 humeri, 1 ulna, 2 coracoids, 1 ischi-um,
partial pubis and ilium, 1 sacral rib, 1 fibula, 3 phalanges, and vertebrae
fragments. An individual specimen consisting of a partial carapace and plastron
missing only the left hypoplastron and xiphiplastron (.299). Isolated skull frag-ment
(parietal, supraoccipital, both maxillae, basisphenoid, 1 quadrate, 1 postor-bital,
1 partial squamosal) (.299.1). A single sub-adult individual with partial
plastron and 2 peripherals (.12 1.1 -.12 1.2).
Isolated elements: 17 nuchals (SCSM 93.90.2-.8)(5 USNM)(5 UF); 3
right and 2 left 2nd costals (.334-.338); 6 right and 5 left 3rd costals (.339-349);
6 left and 6 right 4th costals (.350-.361); 3 right and 2 left 5th costals (.362-366);
7 right and 4 left 6th costals (367-377); 1 left 7th costal (378); 34 peripherals
(300-333); 5 right and 3 left epiplastra (379-386); 3 entoplastra (.417-.419); 5
right and 13 left hyoplastra (387-399,.432)(2 USNM)(2 UF); 3 left hypoplasia
(.400-.402); 6 right and 10 left xiphiplastra (.403-.416)(l USNM)(1 UF); 1
mandible (.757); 3 humeri (.426-.428); 6 femora (.420-.425).
Characters used for identification: Identification of the two most complete spec-imens
is discussed by Bentley and Knight (1993).
Nuchals - See Bentley and Knight (1993), for characters used to distin-guish
this from other possible identifications.
Epiplastron - Differs from C. picta in that the scute overlap area is more
robust in C. guttata, and the anterior edge is not serrated as is common with C.
picta. It can be separated from C. muhlenbergii because the bulbous area where
the gular sulcus wraps onto the scute overlap is usually considerably wider medi-ally
to laterally in specimens of C. guttata, whereas the scute overlap portion that
is posterior to the bulbous area is narrower in C. muhlenbergii than C. guttata.
The length of the scute overlap posterior to the gular sulcus is longer than that of
C. picta. Also, the epidermal attachment scar in C. muhlenbergii is deeply
incised and tends to undercut the scute overlap area. The epiplastron of C. gut-tata
is rarely incised to this extent. The epiplastra of C. insculpta differ from C.
guttata in that the bulbous area where the gular sulcus crosses the dorsal surface
is only slightly or not at all bulbous in specimens of similar size. The epiplas-tron
of Deirochelys, Trachemys and Pseudemys that fall within the size range of
C. guttata have less pronounced scute overlap area compared to that of C. gutta-ta.
E. blandingii specimens of comparable size show sub-adult traits (incomplete
ossification) and have a less pronounced scute overlap area. Terrapene epiplas-tra
differ in that the area posterior to the scute overlap is concave, forming a
depression posteromedially to the gular sulcus, generally absent in C. guttata.
Turtles 1
5
Hyoplastron - Differs from Terrapene and Emydoidea in that C. gutta-ta
lacks the hinge components. C. muhlenbergii differs slightly from C. guttata
in that the scute overlap is narrower in C. muhlenbergii. Specimens of Trache-mys
and Pseudemys that fall within the size range of C. guttata have scute over-laps
that are greatly reduced in comparison to C. guttata and the elements exhib-it
incomplete ossification. Deirochelys and Chrysemys picta can be separated
from Clemmys guttata because the distance between the entoplastron and
hypoplastron is ca. 40% greater in adults of the former two genera. Also, in C.
picta, the humeral sulcus does not cross dorsally over the scute overlap area. C.
insculpta exhibits incomplete ossification when elements fall within the size
range of C. guttata.
Hypoplastron - This element can be separated from other genera of
emydid turtles by the lack of hinge components (separating it from Terrapene
and Emydoidea), or by its posterior width being greater than its length (separates
it from Trachemys and Pseudemys). Holman (1977) gives characters used to sep-arate
this element from C. muhlenbergii and C. insculpta.
Xiphiplastron - Separation of this element from C. picta is listed under
that species. It can be separated from C. muhlenbergii and C. insculpta in that the
posterior edge is generally squared off rather than tapering to a point, as in the
other two species. However, some specimens of C. guttata do exhibit a pointed
condition. These still can be separated from C. muhlenbergii because the area
where the abdominal muscle attaches to the xiphiplastron is more pronounced.
C. insculpta can also be separated from C. guttata because, in the area where the
anal sulcus crosses onto the scute overlap area, the xiphiplastron is deeply
notched, being greatly reduced or lacking in C. guttata.
Entoplastron - In C. guttata the humero-pectoral sulcus crosses the
entoplastron within the anterior half of that element. In C. muhlenbergii the
humero-pectoral sulcus may cross the entoplastron at its posterior extremity, but
typically it does not cross the entoplastron at all (Bentley and Knight 1993).
These fossil entoplastra are tentatively assigned to C. guttata, and not C. insculp-ta,
because this element is generally more robust in C. insculpta and the humero-pectoral
sulcus crosses the entoplastron more posteriorly in C. insculpta than in
C. guttata.
Costal - Characters used to differentiate costal elements from C. picta
are described in that section. Costals of Clemmys guttata differ from C. muh-lenbergii
in having substantially more curvature. The dorsal surface of costals in
C. guttata is smooth, lacking any exterior bulges or sculpturing common to adult
C. muhlenbergii, C. insculpta, Deirochelys, Trachemys, and Pseudemys. The
costals of Emydoidea, Trachemys, and Pseudemys exhibit immature traits when
they are within the size range of C. guttata. Terrapene has deeply incised sulci,
and its elements tend to be more acutely angled proximally and exhibit "bul-bous"
sculpturing.
1 6 Curtis C. Bentley and James L. Knight
Peripherals, femora, humeri, and mandible - These elements are tenta-tively
referred to this species because they compare most closely to Recent and
fossil C guttata.
Remarks: The spotted turtle ranges from northern Illinois into Ohio and Ontario,
east to Maine and New York, and south along the Atlantic Coastal Plain into
northern Florida (Conant and Collins 1991). Clemmys guttata occurs most com-monly
in bogs or marshy pastures, but it also can be found in woodland streams.
It favors habitats with soft substrates. C. guttata is frequently found away from
water, but even so it is the least terrestrial of the three eastern species of Clem-mys
(Ernst and Barbour 1989). The fossil spotted turtle remains from the Ardis
local fauna represent the oldest known material of this species (Bentley and
Knight 1993), and the first fossil record for the eastern United States. Holman
(1990) reports a right epiplastron from a 6,000-year-old fauna near Lansing,
Michigan. Interestingly, Ernst and Barbour (1989) noted a relationship between
this turtle and the burrows of muskrats, which the turtles apparently use for esti-vation
and hibernation sites. Fossil muskrats were the most common mammals
found at the Ardis site (Bentley et al. 1994) and are believed to have used the
solution cavities as burrow sites. This may help to explain the abundance of this
turtle at the Ardis site.
Clemmys muhlenbergii - Bog turtle (Schoepff, 1801)
Material: An individual consisting of a partial carapace (nuchal, 1st and 2nd left
costals and peripherals, 2 peripherals, numerous shell fragments) and plastron
(both epiplastra, and partial hyoplastron) (.429).
Isolated elements: 2 nuchals (.430-.431) ; 2 right epiplastra (.433-.434).
Characters used for identification: C. muhlenbergii fossils were distinguished
from other emydid turtles based on characters listed in previous sections, along
with additional nuchal and sulci characters given by Bentley and Knight (1993)
(Fig.5).
Remarks: The soft bottoms and slow moving waters of swamps, bogs and
marshes are typical aquatic habitats of the bog turtle (Ernst and Barbour 1989),
but this turtle can also be found on land. Clemmys muhlenbergii has a patchy
distribution in the Northeast, and ranges as far south as northern Georgia and
extreme northwestern South Carolina. This disjunct spatial pattern has been
interpreted as suggesting a larger former range (Smith 1957). The fossil evi-dence
from the Ardis site suggests that the species' range extended at least 250
km farther southward during the late Pleistocene.
Turtles 1
7
This is the second report of fossil material of C. muhlenbergii (Holman
1977) and represents the first fossil record from the eastern United States south
of Allegany County, Maryland. This is the first sympatric occurrence of C. muh-lenbergii
and C. guttata in the fossil record.
Fig. 5 Partial fossil carapace of Clemmys muhlenbergii (.429). from the Ardis
local fauna.
3 cm
1 1 1 1
Terrapene Carolina major- Gulf Coast Box turtle (Agassiz, 1857)
Material: An individual male specimen consisting of a partial carapace (SCSM
91.165.1) lacking it's anterior edge (Fig. 6), complete plastron (SCSM 91.165.2),
partial skull (SCSM 91.165.19), 8 cervical vertebrae (SCSM 91. 165. 10-. 17), 1
humerus (SCSM 91.165.5), both scapulo-acromial processes (SCSM 91.165.6-
.7), both coracoids (SCSM 91.165.8-.9), complete pelvic girdle (SCSM
91.165.3), 1 femur (SCSM 91.165.4), 1 sacral rib (SCSM 91.165.18). An indi-vidual
female specimen consisting of a complete carapace (SCSM 91.163.1) and
plastron (SCSM 91.163.2), 2 cervical vertebrae (SCSM 91.163.10-.il), both
scapulo-acromial processes (SCSM 91. 163. 4-. 5), both coracoids (SCSM
91.163.8-.9), both femora (SCSM 91.163.6-.7), and complete pelvic girdle
(SCSM 91.163.3). An individual female specimen consisitng of a complete
carapace and plastron (SCSM 9 1.1 64.1 -.2), 2 cervical vertebrae (SCSM
91.164.9-.10), 1 humerus (SCSM 91.164.8), 1 scapulo-acromial process (SCSM
91.164.4), both coracoids (SCSM 91.164.5-.6), 1 femur (SCSM 91.164.7), com-plete
pelvic girdle (SCSM 91.164.3), and 1 caudal vertebra (SCSM 91.164. 11).
An individual male specimen with complete carapace (SCSM 91. 166. 1) and pos-terior
half of plastron from hinge (SCSM 91.166.2). An individual female spec-
1 8 Curtis C. Bentley and James L. Knight
imen consisting of a partial carapace and one half of posterior plastron from the
bridge (SCSM 91.168.1). An individual female specimen consisting of a com-plete
carapace and plastron (.435-.435.1). A individual partial carapace (SCSM
91. 167.1).
Isolated elements: 8 nuchals (.459-.466); 2 right 1st costals (.471 -.472);
1 fused left 7th and 8th costal (.473); 5 costals (.474-.478); 3 left 5th peripherals
(.479-.481); 2 fused peripherals (1 USNM) (1 UF); 32 peripherals (.510-.541)(2
USNM)(2 UF); 2 complete, 10 partial anterior plastral lobes (.467-.469)(6
USNM)(3 UF); 1 right epiplastron (.470); 2 entoplastra (.508-.509); 2 complete,
9 partial posterior plastral lobes (.507)(5 USNM)(5 UF); 32 large shell fragments
(.510-.541); 1 partial skull (.436); 1 right maxilla (.437); 3 postorbitals (.438-
.440); 4 mandibles (.441-.444); 3 cervical vertebrae (.456-.458); 4 humeri (.445-
.448); 4 femora (.449-.451); 4 ilia (.452-.455).
Characters used for identification: The more complete specimens are easily sep-arated
from other emydid turtles based on their hinged plastra and overall mor-phology.
Emydoidea differs from the box turtle by its smooth, unkeeled carapace
and tends to be anteriorly constricted (Holman 1985).
Plastron - The hinged plastral elements prevent confusion with any
other emydid of North America except E. blandingii. The anterior end of this
attachment area between the carapace and plastron differs from E. blandingii, as
the carapace and plastron of Terrapene have a heavily sutured interlocking pro-trusion
and pocket respectively. In Emydoidea this area lacks the sutured "ball
and socket" mechanism and instead has a pronounced lateral flare generally
located on the 5th marginal. The large size and robust nature of the fossil ele-ments
suggest an affinity to this subspecies.
Peripherals - These elements are distinguished by an upwardly curved
anterior edge, forming, in some specimens, a "gutter-like" effect.
Humeri, femora, and ilia - Humeri and femora were separated from
other emydid turtles based on comparison to Recent specimens and characters
provided by Holman (1967, 1975). Ilia of T. Carolina have a distinctive
"boomerang-shape" and are straighter in other species (Holman 1977). Addi-tionally,
these fossil elements were identical to those retrieved from within the
shells of the more complete fossil T. c. major collected from the Ardis site.
Remarks: This is the largest of the North American box turtles and today ranges
from the coast of eastern Texas eastward along the Gulf coast into the Florida
panhandle (Carr 1952). The Gulf Coast box turtle is commonly found in marsh-es,
palmetto-pine forests, and upland hammocks. It enters water with a frequen-cy
similar to T. c. Carolina (Carr 1952, Conant and Collins 1991).
Turtles 1
9
Fig. 6 Partial fossil carapace of Terrapene Carolina major (SCSM 91.165.1)
which contained the skull (SCSM 91.165.19) shown in figure 6d.
Large fossil box turtle remains have generally been referred to as T. c. putnami
or T. c. putnami x major (Milstead 1969). T. c. putnami differs from T. c. major
only in size, attaining lengths upwards of 300 mm (Auffenberg 1967, Milstead
1969). The largest Recent specimen of T. c. major on record has a carapace
length of 216 mm (Conant and Collins 1991). Auffenberg (1967) reported a
large box turtle (233 mm), with skull, from Haile 8A, stating that it was very sim-ilar
to T. c. major. Blaney (1971) placed T. c. putnami in synomyny with T. c.
major, based on the shared characters of the two subspecies, and stated that size
alone was not a justifiable reason to recognize a subspecies. The fossil box tur-tles
from the Ardis site exhibit all the characters Milstead (1969) used to distin-guish
T. c. putnami. Furthermore, the five specimens had carapace lengths of
190.0 mm to 260.0 mm. Shell and axial elements of fossil box turtles from the
Ardis site could not be consistently distinguished from Recent specimens of T. c.
major except by size. Milstead (1969) stated that populations of T. c. Carolina in
Massachusetts and Michigan, on the northwestern edge of the subspecies range,
appear to have strong morphological affinities to T. c. major having average cara-pace
lengths of 140 mm and 139 mm respectively. Milstead suggested that this
relationship may be due to a pre-Wisconsin influence of T. c. putnami or the
influence of T. c. triunguis. An isolated, fused posterior plastral lobe (82.26 mm)
collected from the Ardis site was estimated to belong to a specimen with a cara-pace
length of about 140-145 mm. This plastral lobe might indicate the presence
20 Curtis C. Bentley and James L. Knight
of box turtles at or near the size of the northwestern populations discussed by
Milstead (1969). Additionally, several significantly smaller isolated peripherals
were collected from the Ardis site, but these peripherals are largely unfused and
may represent juveniles. The association of the smaller fused plastral lobe with
significantly larger specimens suggests that during the height of the Wisconsin
glaciation there may have been intergradation between local and northerly dis-placed
populations of T. c. Carolina, and populations of T. c. major, T. c. triun-guis,
and T. c. baud radiating from the south. The Ardis population, being pre-dominantly
large box turtles, suggests that T. c. major traits (very large size,
elongated shells, and upward curvature of peripherals) were more predominant
during this time period.
The Ardis site produced two partial fossil skulls; SCSM 91.165.19 asso-ciated
with a carapace of 245-250 mm. in length, and (.436), an isolated partial
skull comparable in size to SCSM 91.165.19 (Fig. 6 and Fig. 7). We failed to
distinguish any consistent differences between our skulls and Recent specimens
except for size and an exaggerated upward curvature of the supraoccipital crest
in specimen SCSM 91.165.19. This extreme curvature is considered an anom-aly
due to its complete absence in other fossil specimens; however, it is noted in
Recent specimens but is less developed. The supraoccipital is thought to be one
of the most variable cranial elements for box turtle systematics (W. Auffenberg,
University of Florida, personal communication).
The parietals of most Recent specimens examined of Terrapene Caroli-na
had parietals that were inflated anteriorly, with a reduction in the tabled, dor-sal
surface of this element. Although extremely preliminary, we suggest a cor-relation
between size and the degree of parietal inflation in T. c. major. Anteri-or
inflation of the parietals is greatly reduced to absent among the largest speci-mens.
In other subspecies of T. Carolina, the inflation of the parietals remains
fairly constant. The two skulls from the Ardis site do not exhibit anterior infla-tion
of the parietals. The morphological similarities between the fossil box tur-tles
of the Ardis site and the Recent and fossil specimens examined from muse-um
collections (Fig. 7) suggests that the greatest affinity of the Ardis specimens
is to T. c. major. They support the synonymy of T. c. putnami with T. c. major
(Blaney 1971). Affinities noted by Milstead (1969) between northwestern pop-ulations
and T. c. major may be a result of the proposed intergradation between
box turtles during the height of the Wisconsin glaciation. The synonymy of T. c.
major with T. c. putnami also suggests that T. c. major may have been capable of
obtaining a considerably larger size than that observed in living specimens.
Although we believe a systematic revision of the genus Terrapene is
needed, it exceeds the bounds of this faunal review. One of the goals of this dis-cussion,
however, is to emphasize the need for such a revision, based both on
"standard" characters and on osteological characters of all fossil and extant
forms.
Turtles 21
Fig. 7 Terrapene Carolina major skulls. A) Recent (UF 18963). B) Haile 8A (UF
3148). C) Ardis fossil (SCSM 91.165.19). D) Ardis fossil (.436).
3 cm
3 cm B
3 cm D
22 Curtis C. Bentley and James L. Knight
Deirochelys reticularia - Chicken Turtle (Agassiz, 1857)
Material: 2 peripherals (.542-.543).
Characters used for identification: Identification is based upon the distinctive
"spike-like" pattern of the dorsal sculpturing on these elements (Jackson 1964,
Holman 1978).
Remarks: This turtle today has a continuous range from Texas and Oklahoma
through the Gulf states and along the southern half of the Atlantic Coastal Plain
states into North Carolina, with isolated populations in southeastern Virginia
(Conant and Collins 1991). Still-water habitats, such as ponds, swamps, marsh-es,
and temporary pools, are commonly occupied by chicken turtles, which
reportedly do not favor moving water (Ernst and Barbour 1989).
This is the first fossil record of Deirochelys from South Carolina. The
species most extensive fossil record and probable origin is in Florida (Jackson 1978).
Emydoidea blandingii - Blanding's Turtle (Holbrook, 1838)
Material: An individual specimen consisting of a complete carapace, anterior
plastral lobe (.544), left lower jaw (.544.1), partial hyoid (.544.2), 1 partial
humerus (.544.4), 1 partial scapulo-acromial process (.544.3), 1 femur (.544.5),
3 partial dorsal vertebrae (.544.8-.544. 10), 1 ilium (.544.6), 1 pubo-pectineal
process (.544.7), 1 sacral rib (.544.11). An individual specimen consisting of a
complete carapace, 1 phalange, 2 partial vertebrae (UF). An individual specimen
consisting of a nearly complete carapace, anterior plastral lobe, 4 partial dorsal
vertebrae, 3 caudal vertebrae (USNM). An individual specimen consisting of a
complete carapace, posterior plastral lobe (.545), 5 cervical vertebrae (.545.1-
.545.5), 2 humeri (.545.21-.545.22), 1 ulna (.545.31), 2 scapulo-acromial
processes (.545.27- .545.28), 2 coracoids (.545.25-.545.26), 1 dorsal vertebra
(.545.6), 2 femora (.545.23-.545.24), 1 fibula (.545.29), 1 tibia (.545.30), 1 com-plete
pelvic girdle (.545.20), 5 phalanges (.545.32-.545.35), 1 ungual (.545.36),
2 sacral ribs (.545.37-.545. 38), 13 caudal vertebrae (.545.7-.545. 19). An indi-vidual
specimen consisting of a complete carapace and plastron, partial skull (2
maxillae, 1 quadrate, basioccipital-condyle, basisphenoid, frontal-postorbital-parietal
skull fragment) (.546), partial hyoid apparatus (.546.2), 6 cervical verte-brae
C546.3-.546.8), 1 humerus (.546.20); 2 ulnae (.546.25-.546.26), 1 radius
(.546.27), 2 scapulo-acromial processes (.546.28-.546.29), 2 coracoids (546.30-
.546.31), 1 femur (.546.21), 1 tibia (.546.22), 2 fibulae (.546.23-.546.24), com-plete
pelvic girdle (.546.19), 10 phalanges (.546.32-.546.41), 1 ungual (.546.42),
2 sacral ribs (.546.43), 10 caudal vertebrae (.546.9-.546. 18). An individual spec-imen
consisting of a complete carapace (.547). An individual specimen with the
Turtles 2 3
anterior one half of the carapace (.548). An individual specimen with the ante-rior
two-thirds of the carapace (.549). An individual juvenile specimen consist-ing
of a partial carapace (nuchal, 1-2,4,8 neurals, pygal, 2-5, 7-11 left peripher-als,
1-4 right peripherals, 3-8 right costals, 3rd left costal) and anterior lobe of
plastron missing left epiplastron (.550), left xiphiplastron (.550.1), 1 dorsal ver-tebra
(.550.2).
Isolated elements: 1 nuchal (.588); associated nuchal and 1st left and
right costal with 1st left peripheral (USNM), 4 associated costals and single
peripheral (.566); 1 left and 1 right 1st costal (.567-.568); 1 left 3rd costal (with
rodent gnaw marks) (.569); 10 costals (.556-.565); 6 neurals (.570-.575); 3 right
1st peripherals (.576-.578); 1 left 5th peripheral (sub-adult) (.580); 1 left 6th
peripheral (.579); 1 partial plastron (.607); 2 anterior plastral lobes (2 UF), 1 par-tial
anterior plastral lobe (USNM), 2 pairs of associated epiplastra (.589-. 590); 4
left and 2 right epiplastra (3 USNM)(3 UF), 7 entoplastra (.581-.587); 4 left and
6 right hyoplastra (.601-.602)(4 USNM)(4 UF); 4 posterior plastral lobes
(.591)(1 USNM)(2 UF); 1 associated right hypoplastron and xiphiplastron
(.592); 7 right and 8 left hypoplastra (.593-.600)(3 USNM)(4 UF); 3 right and 3
left xiphiplastra (.603-.606)(l USNM)(1 UF); 2 partial skulls (.551-.552); 1 left
postorbital (.553); 2 left auditory bullae and quadrate (.554-.555); 5 cervical ver-tebrae
(.614-.618); 3 humeri (.608-.610); 3 femora (.611-.613); 1 partial pelvic
girdle (.619).
Characters used for identification: It is possible to distinguish the more complete
specimens of Emydoidea blandingii from Deirochelys reticularia on the basis of
the hinged plastral elements and the lack of carapacial sculpturing in the former
(Jackson 1978). Characters that distinguish this species from Terrapene Carolina
are given under that account. Isolated specimens can be distinguished from other
emydid turtles based on characters mentioned in other sections of our paper and
the following:
Epiplastron - Emydoidea epiplastron can be contrasted to Terrapene
epiplastron in several ways. This element in Emydoidea differs from Terrapene
by the presence of a depression located on the dorsal surface and medially to the
anterior scute overlap area, which is less pronounced in Emydoidea. When com-pared
to specimens of Terrapene of comparable size, this element is less robust
and somewhat dorso-ventrally compressed. However, there is some difficulty in
distinguishing large specimens of T Carolina major from E. blandingii. This ele-ment
differs from Trachemys in that it is more elongated and thinner in E.
blandingii.
Xiphiplastron - This element is most easily confused with Clemmys.
Preston and McCoy (1971), suggested that the xiphiplastron of Clemmys is wider
with respect to its length. Preston (1979) discusses additional characters used to
identify this element in Emydoidea.
24 Curtis C. Bentley and James L. Knight
Neurals - These thin, very broad, smooth elements are distinctive
among emydid turtles.
Axial and appendicular skeleton - Although cranial material of this
species is distinctive, the generalized nature of the postcranial material makes the
assignment to E. blandingii tentative.
Remarks: The recent distribution of E. blandingii is limited to southern Ontario
and the Great Lakes region, with scattered populations occuring westward into
northeastern Nebraska and south into northeastern Missouri, and eastward into
New York and Massachusetts on the Atlantic coast (Conant and Collins 1991).
The nearest fossil records of Emydoidea blandingii to the Ardis site are from
Catalpa Creek, Mississippi (Jackson and Kaye 1975), and at New Trout Cave,
West Virginia (Holman and Grady 1987). Both records are late Pleistocene.
The well preserved fossil material from the Ardis site (Fig. 8) is the first
report of this species on the Atlantic Coastal Plain and is a range extension of
about 1,200 km from its present continuous distribution and nearly 525 km south
of the nearest reported fossil locality at New Trout Cave.
Habitats frequented by E. blandingii are generally in shallow, lentic
waters with soft substrate, such as ponds, streams, marshes and sloughs (Ernst
and Barbour 1989).
Fig. 8 Fossil Emydoidea blandingii carapace (.547).
5 cm
Turtles 2 5
Trachemys scripta - Slider Turtle (Schoepf, 1792)
Material: SCSM 91.169.1 a nearly complete carapace and plastron, partial right
lower jaw, 1 scapulo-acromial process, 1 coracoid, 1 partial vertebra, partial
pelvic girdle, 1 fibula. An individual specimen consisting of a partial carapace
and plastron (.620).
Isolated elements; 3 nuchals (.621 -.623); 23 partial costals (.624-
.640)(3 USNM)(3 UF); 15 peripherals (.641-.655); 1 suprapygal (.656); 10 neu-rals
C657-.666).
Characters used for identification: This turtle is distinguished from other turtles
by characters given by Holman (1985). Additionally, it has a diagnostic carapa-cial
ornamentation.
Remarks: The slider has a nearly continuous distribution from Illinois southward
into Texas and New Mexico and thence eastward across northern Florida north
along the Atlantic coast into Virginia, with populations in Mexico and Maryland
(Conant and Collins 1991). Trachemys scripta can be found in most freshwater
habitats, but seems to prefer slow to non-moving water with a soft substrate
(Ernst and Barbour 1989).
Today Trachemys scripta is common around the Ardis site and we have
observed more than 10 within 100 m of the excavation site.
Pseudemys sp.-Cooters (Gray, 1855)
Material: 1 right 1st peripheral (.667); associated 9- 10th right peripherals (.668);
2 peripherals (.669-.670).
Characters used for identification: These compare most favorably to species in
this genus based on the thin, elongated sloping nature of the elements. We were
unable to identify any diagnostic characters on these elements that could be used
make a species placement. The only genus with which these may be easily con-fused
is Trachemys. The fossil elements lack the sculpturing found in Trache-mys
and are significantly thinner and more elongated. The fossil peripherals also
have a straight distal margin which contrasts with the notched margin in Trache-mys.
26 Curtis C. Bentley and James L. Knight
Remarks: Pseudemys is a common genus of the Southeast, found in various
aquatic systems (Conant and Collins 1991). The two species found in South Car-olina
today are P. floridana and P. concinna.
Testudinidae
Hesperotestudo crassiscutata -Giant Tortoise (Williams, 1950)
Material: 9 carapacial and plastral fragments (.671-.677)(1 USNM)(1 UF); 1
right xiphiplastron (.678); 1 vertebra (.685); 1 ungual (.686); 6 osteoderms (.679-
.684).
Characters used for identification: The fragmented shell elements were assigned
to this species rather than H. incisa on the basis of their extremely large size and
robustness (40.0 mm thick). The large osteoderms, phalange, and vertebra could
not be distinguished from H. crassiscutata in the Florida Museum of Natural
History.
Remarks: Bramble (1971), Preston (1979), and Meylan (1995) place all North
American non-Gopherus tortoises into the genus Hesterotestudo, and that prac-tice
is followed here. Dobie and Jackson (1979), provided the first report of
(Geochelone) H. crassiscutata in South Carolina from the late Pleistocene of
Edisto Island.
Trionychidae
Apalone sp.- Softshell turtle (Rafinesque, 1832)
Material: 1 partial nuchal (.47); 1 costal distal end (.48); 1 partial neural (.49).
Characters used for identification: These fossils are easily assigned to this genus,
based on the relatively thin shell elements with characteristic pitting of the dor-sal
surfaces, general morphology, and the geographical distribution of Triony-chidae.
However, based on these elements, we were unable to identify a species
with any certainty.
Remarks: Two species of the genus Apalone now occur in South Carolina, A.
ferox and A. spinifera. Both species inhabit various aquatic environments with
muddy or sandy bottoms in deep or shallow water (Ernst and Barbour 1989).
Apalone ferox occurs throughout Florida and in the southern portions of Alaba-ma,
Georgia, and South Carolina. A. spinifera is restricted in Florida to rivers in
the extreme northeastern and northwestern portions, and ranges no farther north
along the Atlantic coast than North Carolina. It ranges westward into Colorado
and north into Minnesota, with isolated populations in Montana, California, and
Turtles 2 7
Mexico (Conant and Collins 1991). Only A. spinifera inhabits the area of the
Ardis site today. Dobie and Jackson (1979) reported "Trionyx sp." from Edisto
Island, South Carolina. Meylan (1987) has shown the correct name for North
American softshells to be Apalone.
DISCUSSION AND PALEOECOLOGY
The turtle assemblage of the Ardis local fauna provides important new
data for the late Pleistocene of the southeastern United States. In particular, it
documents a shift in the spatial patterns of several turtle species during the late
Pleistocene. The turtle fauna is unique in its geographical and temporal setting,
and it contains the first sympatric fossil occurrences of several taxa, e.g., Clem-mys
muhlenbergii, C. guttata, Emydoidea blandingii, Macroclemys temminckii.
All of the fossil turtles collected from the site, except Hesperotestudo
crassiscutata and Terrapene Carolina major, are primarily aquatic and are com-monly
found in, or require, still or slow moving water with a soft substrate and
aquatic vegetation (Ernst and Barbour 1989). Additional evidence of a nearby
body of water included the presence of Alligator mississippiensis and elements
of the fish fauna which are currently under study. This agrees with the habitat
suggested by Bentley et al. (1994) based on the Ardis mammal fauna that indi-cates
an ecotone between a mixed forest of conifers, hardwoods and meadows,
and a permanent body of water such as a river or stream which may have given
way to a bog or marsh. Portions of the mammalian fauna and avian material
from the Ardis site further suggest the presence of a nearby large body of water
such as a lake or pond. This association is based on the life histories and habitat
requirements of many of the extant species represented in the Ardis fauna.
The Ardis turtle fauna consists of thirteen extant and one extinct
species, with five taxa considered extralimital. Three of the five have northern
affinities: Emydoidea blandingii, Clemmys muhlenbergii, and Chrysemys picta.
Terrapene Carolina major has a strong southern affinity. Macroclemys has a pri-marily
Gulf coast distribution but extends as far north up the Mississippi Valley
as Iowa (Pritchard 1989). Although Hesperotestudo was widespread in North
America by the Miocene, Hibbard (1960) suggested that the presence of Hes-perotestudo
crassiscutata in a fauna indicated a mild climate with frost-free win-ters.
The sympatric occurrence of species that are apparently ecologically
incompatible today constitutes a "disharmonious fauna" (sensu Lundelius et al.
1983), which has been interpreted by many authors (Hibbard 1960, Holman
1980, Lindelius et al. 1983) as indicating a more equable climate (reduced sea-sonal
temperature gradients) than that experienced in the region today. The
Ardis local fauna reflects such a disharmonious biota, which clearly has no mod-ern
analogue. The turtle fauna corroborates conclusions made on the basis of the
Ardis mammal fauna (Bentley et al. 1994), which also suggests that a more
28 Curtis C. Bentley and James L. Knight
equable climate than today prevailed near or during the maximum advance of the
Laurentide ice sheet in the southeastern United States.
Based on the distribution of terrestrial vertebrates, Smith (1957) stated
that the northeastern biota of North America was displaced southward during the
Wisconsinan glacial maximum. Smith also suggested that during a "Climatic
Optimum," southern counterparts dispersed northward. During the height of the
Wisconsinan glaciation, E. blandingii would have been extirpated from its pre-sent-
day northerly distribution (Mickelson et al. 1983, Conant and Collins
1991). The southern boundary of the Laurentide ice sheet, while abutted against
the Appalachian Plateau (Mickelson et al. 1983), could potentially have forced
the range of E. blandingii to be split, with one population occurring along the
Mississippi Valley and another along the Atlantic Coastal Plain. This may have
produced geographically isolated eastern and western populations of Emydoidea
blandingii during the late Pleistocene.
In addition to Emydoidea blandingii, Spermophilus tridecemlineatus
(thirteen-lined ground squirrel) was also collected from the Ardis site (Bentley et
al. 1994). Based on Recent and fossil distributions (Kurten and Anderson 1980),
the present northeastern distribution of the thirteen-lined ground squirrel also
may have been displaced southward along the Atlantic Coastal Plain by the
advancing Laurentide ice sheet.
As with the Gulf Coast Corridor, depressed sea levels during the Pleis-tocene
glacial stage exposed much or all of the Atlantic continental shelf (Bloom
1983), thereby widening the Atlantic Coastal Plain. This may have facilitated
the dispersal of glacially-displaced species southward along the coast. The
newly emergent land area provided expanded habitats for species to utilize
(Blaney 1971), and a more equable climate may have allowed for the range
extention of both northern and southern species into these new areas. Northern
populations of T. c. Carolina in Massachusetts and Michigan that show affinity to
T. c. major, discussed by Milstead (1969), may be relicts left over from a Pleis-tocene
interval of extensive intergradation between the northerly displaced sub-species
and radiating southern subspecies.
Bleakney (1958) suggested that the Recent population of E. blandingii
in Nova Scotia survived glaciation in an "Atlantic Coastal Plain refuge" and then
dispersed northward up the coast into Canada and Maine. Preston and McCoy
(1971), suggested that "colonization of the Atlantic Coastal Plain from the Great
Lakes region, along a 'steppe corridor' (Schmidt 1938) through the Mohawk
Valley" was a more plausible hypothesis. Preston and McCoy (1971) also stat-ed
that the study of specimens from the eastern limits may provide an answer to
the possibility of a "minor Atlantic Coast refuge for Emydoidea during ice
advances." The presence of numerous E. blandingii at the Ardis site, as well as
fossil material from New Trout Cave in West Virginia (Holman and Grady 1987),
suggest that the Recent extreme northeastern populations may be products of a
Turtles 2 9
re-invasion of the northeast by a substantial Atlantic Coastal Plain stock that had
spread at least 900 km southward during the glacial advance of the late Pleis-tocene.
Evidence from the Ardis local fauna indicates significant shifts longitu-dinally
in the spatial distribution of E. blandingii along the Atlantic Coastal
Plain during the late Pleistocene.
ACKNOWLEDGMENTS - We wish to express our gratitude to the staff of the
Giant Cement Plant, Harleyville, South Carolina, for their generous cooperation,
and in particular to Burt Ardis, for whom the fauna is named. Many thanks go
to all the field volunteers: Vance McCollum, Linda Eberle, Craig and Alice
Healy, Derwin Hudson, Ray Ogilvie, Lee Hudson, Tom Reeves, Martha Bentley,
and Karin Knight. Robert Weems, U.S. Geological Survey, and Peter Meylan,
Eckerd College, Petersburg, Florida, provided helpful discussion and review of
this manuscript. We also wish to express our appreciation to Overton Ganong of
the South Carolina State Museum; David Webb, Gary Morgan, and David Auth,
of the Florida Museum of Natural History; Dennis Hermann, of Zoo Atlanta; and
Greg Schneider, Museum of Zoology, The University of Michigan, and Justin
Cougdon, Savannah River Ecology Labratory, Aiken, South Carolina, for allow-ing
us access to their collections. Thanks go to Mike and Debi Trinkley, the
Chicora Foundation, Columbia, South Carolina, for the use of their field equip-ment,
and to Mike Runyon of Lander College, Greenwood, South Carolina, for
donating fossil material for C14 dating. Darby Erd provided the illustrations.
LITERATURE CITED
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1
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32 Curtis C. Bentley and James L. Knight
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Testudinidae) based on new fossil records from Kansas and Oklahoma.
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Received 3 January 1996
Accepted 13 August 1996
Turtles 3 3
APPENDIX I
Taxa and Minimum Number of Individuals Present
Taxa Minimum number of individuals
Kinosternon subrubrum* 5
Sternotherus odoratus* 4
Chelydra serpentina 3
Macroclemys temminckii* 1
Clemmys guttata 19
Clemmys muhlenbergii* 3
Chrysemys picta* 25
Deirochelys reticularia* 1
Trachemys scripta 4
Pseudemys sp. 1
Terrapene Carolina major 12
Emydoidea blandingii* 16
Hesperotestudo crassiscutata 1
Apalone sp. 1
Number of turtle species = 14
Number of individuals = 96
* = first fossil report from South Carolina
Observations of Freshwater Jellyfish, Craspedacusta sowerbyi
Lankester (Trachylina: Petasidae), in a West Virginia Reservoir
Ted R. Angradi
U.S. Department ofAgriculture, Forest Service
Timber and Watershed Laboratory,
Parsons, West Virginia 26287
ABSTRACT. — A swarm of medusae of the freshwater jellyfish
Craspedacusta sowerbyi was observed in a cove of a West Virginia
reservoir in August and September, 1995. Medusae were abundant
(>1000/iTr) but extremely localized. Distribution of medusae in the
cove did not appear to be linked to water chemistry. Size of medusae
ranged from 6-21 mm in diameter and increased significantly with dis-tance
from the center of abundance, suggesting that the localized distri-bution
of medusae resulted from dispersion rather than from environ-mentally-
induced aggregation. Measurements of mean diameter of
medusae on separate dates indicated a growth rate of about 0.2 mm/d,
and a medusa life cycle of approximately 102 days.
The freshwater jellyfish Craspedacusta sowerbyi Lankester 1880 is an
exotic species first observed (as medusae) in the United States in 1908 (Kramp
1950, Pennak 1989). Native to the Yang-tse River system in China (Kramp
1950), C sowerbyi has been reported from many localities worldwide between
45° north and 45° south latitude (Acker and Muscat 1976, Pennak 1989). True
freshwater jellyfishes are few, limited to about a dozen species worldwide
(Hutchinson 1967, Pennak 1989).
Craspedacusta sowerbyi has been reported from 3 1 states and the Dis-trict
of Columbia; it has not been reported from northern New England, the
Northern Rocky Mountains, or the Northern Great Plains (DeVries 1992). In
West Virginia there are records of C. sowerbyi from Barbour, Fayette, Mercer,
Monogalia, Wayne, and Wood counties (Reese 1940, Lytle 1962, Koryak and
Stafford 1981, and D. Tarter, Marshall University, personal communication).
Craspedacusta sowerbyi has two life stages, a free-swimming medusa
(10-20 mm diameter), and a sessile hydroid polyp (1 mm long, Acker and Mus-cat
1976). Lytle (1959) reviewed the developmental biology of the species.
Medusae of C. sowerbyi appear sporadically in lentic and even less frequently in
lotic ecosystems in the United States (Acker and Muscat 1976, Beckett and
Turanchik 1980, DeVries 1992). Usually a swarm of medusae appears in sum-
34
Jellyfish 35
mer where it has never before been observed or where it has not been observed
for many years (Slobodkin and Bossert 1991). Lentic systems in which the
medusae have been observed include reservoirs, natural lakes, ponds, quarries,
ornamental pools, and aquaria.
Specific environmental factors associated with the formation of
medusae from polyps via asexual reproduction (budding) are poorly understood.
Factors suggested include increasing water temperature (McClary 1959)
increased alkalinity (Koryak and Clancy 1981, McCullough et al. 1981),
increasing dissolved CO2 (Acker and Muscat 1976), decreasing stream flow
(Brussock et al. 1985), changing reservoir levels (Deacon and Haskell 1967) and
increasing supply of zooplankton (Lytle 1959), on which the medusae prey.
Dispersal of C. sowerbyi among water bodies probably occurs via
polyps attached to aquatic plants or waterfowl, or in tanks used to transport fish
(Byers 1945, Bushnell and Porter 1967, Howmiller and Ludwig 1970). The
polyps can survive in moving water (Hutchinson 1967), so once the polyps enter
a river system, the medusae may eventually appear in downstream reservoirs
(e.g., Yeager 1987).
Because the medusae occur unpredictably and the polyps are micro-scopic
and easily overlooked, the complete geographic distribution and ecology
of C. sowerbyi are not well known. Field studies have been mostly descriptive
(e.g., Garman 1916, Deevy and Brooks 1943, Dexter et al. 1949, Chadwick and
Houston 1953, Bushnell and Porter 1967, Koryak and Clancy 1981, McCullough
et al. 1981, Dodds and Hall 1984). Deacon and Haskell (1967) examined diel
activity patterns of medusae at Lake Mead, Nevada. Dodson and Cooper (1983)
examined trophic relationships of the medusae in the laboratory. Acker and
Muscat (1976) and DeVries (1992) reviewed the literature on the ecology of C.
sowerbyi.
The purpose of this paper is to describe a swarm of C. sowerbyi
medusae I observed at Stonewall Jackson Lake, Lewis County, West Virginia, in
August and September 1995. My initial observations of the medusae at
Stonewall Jackson Lake suggested that the size distribution of medusae varied
with distance from the main concentration of medusae (swarm). I hypothesized
that the distribution of medusae resulted from dispersion from the apparent pop-ulation
center at the swarm, and I predicted that medusae collected away from
the main swarm location would be larger than medusae within the swarm
because more distant medusae would have had more time to grow. The null
hypothesis that medusae collected from all locations have the same size class dis-tribution
implies that the dense concentration of medusae at the swarm location
results primarily from aggregation due to water chemistry, temperature, food,
current, wind, or some other factor rather than from dispersion. I collected and
measured specimens to test this hypothesis. I also compared the mean size of
36 Ted R. Angradi
specimens collected on separate dates to calculate an approximate growth rate
for the medusae.
METHODS
I observed the medusae at Wolf Fork (80°28'W, 38°59'N), a cove
formed by a flooded tributary to Skin Creek which forms the east arm of
Stonewall Jackson Lake (Fig. 1). Wolf Fork is about 1.5-km long and 30-150-m
wide and has abundant flooded timber. The cove is well sheltered from winds
and is a no-wake boating zone. A culvert connects Wolf Fork and the stream
draining the upper watershed. Stonewall Jackson Lake is a 1,070-ha Army
Corps of Engineers reservoir filled in 1986. The main West Fork River arm of
the reservoir is a tributary of the Tygart River in the Ohio River drainage.
During my initial visit to Wolf Fork (16 August 1995) I estimated the
density (number m3
) of medusae using two methods. Where the medusae were
abundant I used a 20-L plastic bucket. From a small boat I slipped the bucket
into the water and withdrew it with minimal turbulence. I poured the bucket con-tents
through a fine sieve and transferred the medusae to a tray for enumeration.
Where the medusa were scarce, I estimated the density visually. Both methods
are biased toward the upper 0.5 m of water surface because I could not see or
sample medusa at greater depths. On the first visit to Wolf Fork I collected water
samples and recorded water temperature and dissolved oxygen at several loca-tions
in the cove. Water samples were analyzed at the U.S. Forest Service Tim-ber
and Watershed Laboratory, Parsons, West Virginia.
Fig. 1 Map of Stonewall Jackson Lake showing the swarm location at Wolf
Fork. Inset map shows location of the reservoir in West Virginia.
Jellyfish 37
On subsequent visits to Wolf Fork (16 August and 12 September 1995)
I collected medusae with an aquarium dip net from a small boat. I measured bell
diameter of live specimens under a dissecting microscope by placing a ruler
under a clear plastic petri plate containing a few medusae and a small amount of
water. I judged measurement error to be ±1.0 mm.
RESULTS
On 16 August 1995, I observed a dense swarm of medusae near the
head of Wolf Fork. Medusae decreased greatly in abundance with distance from
the swarm. Density of medusae in three bucket samples was 1.2, 1.9, and 4.8
medusa/L. This is approximately equivalent to 1,000-5,000 medusae/m3 in the
upper 0.5 m of the water column within an area of about 25 m2
. At 100-200 m
from the swarm (toward the main channel of the reservoir) there were 10-50
medusae/m3
; at 300-400 m medusae were scarce (<1 medusa/m3
). I did not
observe medusae in lower Wolf Fork, the main channel, or in other coves of Skin
Creek, although I did not make an exhaustive search. My conversations with
anglers, reservoir managers, and local fish biologists suggest that this is the first
record of C. sowerbyi at Stonewall Jackson Lake. The origin of C. sowerbyi in
the drainage is unknown.
Table 1. Selected water quality measurements for Wolf Fork, 16 August 1995.
Site distances are from the swarm toward the main channel. All values for sam-ples
collected at the water surface. DO=dissolved oxygen.
Site Temp DO PH Conductivity Alkalinity so4 Ca Medusa
(m) C (mg/L) nSlcm mg/L
(CaCO,)
(mg/L) (mg/L) abundance
(#/m<)
-100 30.0 7.7 7.0 98.3 19.1 21.9 12.1 <1
30.0 8.1 7.0 100.5 18.
2
21.6 12.8 1000-5000
100 30.5 8.3 7.1 99.8 17.8 22.0 12.5 10-50
200 30.5 8.3 7.2 99.6 17.5 23.7 12.6 <1
300 30.5 8.3 7.2 98.1 17.2 22.6 12.7 <1
Water chemistry did not differ appreciably among sites within Wolf
Fork (Table 1). Dissolved oxygen decreased with depth to 5 mg/L at 2.1 m and
1.8 mg/L at 2.7 m. Depth of the channel in Wolf Fork was 2.5-3.5 m. Flow from
the Wolf Fork watershed (the stream) into Wolf Fork (the cove) could not be
detected at the culvert.
On 18 August 1995, medusae collected ranged in size from 6 to 20 mm
(Fig. 2). Mean (SE) size and size class distribution of medusae collected from
the swarm location were different from medusae collected at three other stations
in Wolf Fork. Swarm medusae were 13.3 (0.2) mm in diameter; medusae from
outside the swarm were 15.2 (0.3) to 16 (0.4) mm in diameter depending on col
38 Ted R. Angradi
Fig. 2 Size distribution of C. sowerbyi medusae collected 0, 100, 200, and 350+
m from a dense concentration (swarm) of medusae at Wolf Fork, Stonewall Jack-son
Lake, West Virginia on 18 August 1995. Figure legends give distance from
the main swarm location, mean (SE) medusa diameter and sample size.
T3
0)
T5
oo
<D
CO
C/>
13 D
<D
E
CD
.a
ou -
200 m
E3
x=15.8(0.3)
C 20 n=110
16
o
10
ii.l.iJlillllli..
30
20
10
350+ m
x=16.0(0.4)
n=52
4 6 8 10 12 14 16 18 20 22
Medusa diameter (mm)
Jellyfish 39
lection site, a significant difference (Kruskal-Wallis ANOVA on Ranks; H =
75.1, P < 0.01). Among sites outside the swarm location, there was also a trend
of increasing size with distance from the swarm; mean medusa diameter was
largest 350 m from the swarm, the location at which medusae were first observed
when entering Wolf Fork from the reservoir. Within the swarm there was a
bimodal size distribution of medusae concentrated in 9 - 13 mm and 15-18 mm
size classes, with more medusae in the smaller size classes. Away from the
swarm most medusae were in the larger size classes, but a semblance of the
bimodal pattern was apparent (Fig. 2).
Fig. 3 Size distribution of C. sowerbyi medusae collected and 100 m from a
dense concentration (swarm) of medusa at Wolf Fork, Stonewall Jackson Lake,
West Virginia on 18 August and 12 September 1995. Otherwise as for Fig. 2.
18 August 1995
4 6 8 10 12 14 16 18 20 22 4 6 8 10 12 14 16 18 20 22
Medusa diameter (mm)
On September 12, I observed fewer medusae than on previous visits to
Wolf Fork. Water temperature was 25 C, and reservoir level had dropped about
1 m since the previous visit. The sky was overcast and light rain was falling,
whereas the sun shone on previous visits. Medusae were most abundant near the
18 August swarm site; at 100 m they were scarce, and were not observed else-where.
Mean diameter of medusae was the same at both locations (Fig. 3),
although more medusae from the 100 m site were in the largest size classes (20-
21 mm). Between August 18 and September 12, mean diameter of medusae at
the swarm location increased slightly more than 4 mm (Fig. 3). This is equiva-
40 Ted R. Angradi
lent to an average growth rate of about 0.2 mm/d. Assuming the medusa stage
originates at about 0.5 mm and attains a maximum size of about 21 mm (Pennak
1989), this rate of growth indicates a life cycle for the medusa of about 102 days.
The apparently slower growth of medusae away from the main swarm (Fig. 3)
may be attributed to disproportionate mortality of the full-growth medusae.
DISCUSSION
I observed great variation in abundance of medusae over a scale of a
few meters that is not readily explained by available water chemistry data. My
finding of larger individuals away from the main concentration of medusae is cir-cumstantial
evidence that spatial variation in abundance of the medusae results
from dispersion from a highly localized swarm location.
If the polyps are lotic (Hutchinson 1967), then the distribution of the
medusae in Wolf Fork results from events and conditions since delivery of imma-ture
medusae from upstream earlier in summer. During most of the year, flow
from the stream would induce some current at the head of Wolf Fork, and the
location and localization of the swarm at Wolf Fork may have resulted from
flow-related concentration (e.g., in an eddy) of polyps or small medusae origi-nally
exported from the stream.
Several authors have also found a highly localized distribution of
medusae. Dodson and Cooper (1983) found medusae only in one small sheltered
cove of a Wisconsin Lake. Deacon and Haskell (1967) reported medusae from
sheltered coves at Lake Mead, Nevada, and not from the open waters of the reser-voir.
Garman's (1916) description of the location of a dense swarm in a narrow
flooded tributary of an impounded reach of the Kentucky River is similar to Wolf
Fork and to other published accounts (e.g., Lytle 1962).
Few authors have explicitly commented on factors accounting for spa-tial
variation in medusae abundance within a reservoir or lake. Acker and Mus-cat
(1976) noticed an apparent effect of light on the distribution of medusae
which they attributed to a direct light effect or to an indirect effect of light on
food concentration. Other factors such as wind (Deevy and Brooks 1943), and
ebbulition (Koryak and Stafford 1981) have been proposed. I noticed no appar-ent
light effect at Wolf Fork, and the cove is well protected from wind and boat-wake
disturbance; these factors are unsatisfactory for explaining the large local
variation in abundance of medusae at Wolf Fork. My observations suggest
instead that the distribution of medusae may result from the pattern of dispersion
from a location that is determined by the habitat requirements of the polyps in
the reservoir or in tributary streams.
My observation of a bi-modal size distribution of medusae (Fig. 2) sug-gest
the possibility of more than one period of medusa formation at Wolf Creek,
perhaps associated with variation in water temperature, water chemistry, or
streamflow during early summer. McClary (1959) reported medusa budding
Jellyfish 41
only within a narrow temperature range, 26 C to 33 C. However, I found very
few small (<8 mm) medusae in August while sampling within this temperature
range (Fig. 2) suggesting that formation of medusae was not ongoing and most
likely occurred during two periods in early summer.
ACKNOWLEDGMENTS
Cliff Brown of the West Virginia Division of Natural Resources pro-vided
access to Wolf Fork. Emmett Fox conducted water quality analysis. Neal
Auvil brought the jellyfish to my attention. Linda Plaugher assisted with the fig-ures.
LITERATURE CITED
Acker, T S., and A. M. Muscat. 1976. The ecology of Craspedacusta sowerbii
Lankester, a freshwater hydrozoan. The American Midland Naturalist
95:323-336.
Beckett, D. C, and E. J. Turanchik. 1980. Occurrence of the freshwater jelly-fish
Craspedacusta sowerbyi Lankester in the Ohio River. Ohio Journal of
Science 80:95- 96.
Brussock, P. P., L. D. Willis, and A. V. Brown. 1985. Flow reduction may
explain sporadic occurrence of Craspedacusta sowerbyi (Trachylina)
medusae. Proceedings of the Arkansas Academy of Science 39: 120.
Bushnell, J. H., and T W. Porter. 1967. The occurrence, habitat, and prey of
Craspedacusta sowerbyi (particularly polyp stage) in Michigan. Transac-tions
of the American Microscopy Society 86:22-27.
Byers, C. F. 1945. The fresh-water jellyfish in Florida. Proceedings of the Flori-da
Academy of Sciences 7:173-180.
Chadwick, C. S., and H. Houston. 1953. A "bloom" of fresh water medusae
Craspedacusta ryderi (Potts) in Kentucky Lake, Tennessee. Journal of the
Tennessee Academy 28:36-37.
Deacon, J. E., and W. L. Haskell. 1967. Observations on the ecology of the
freshwater jellyfish in Lake Mead, Nevada. The American Midland Natu-ralist
78:155-166.
Deevey, E. S., and J. L. Brooks. 1943. Craspedacusta in open water, Lake
Quassapaug, Connecticut. Ecology 24:266-267.
DeVries, D. R. 1992. The freshwater jellyfish Craspedacusta sowerbyi: a sum-mary
of its life history, ecology and distribution. Journal of Freshwater
Ecology 7:7-16.
Dexter, R. W., T. C. Surrarrer, and C. W. Davis. 1949. Some recent records of
the freshwater jellyfish Craspedacusta sowerbii from Ohio and Pennsylva-nia.
The Ohio Journal of Science 49:235-241.
Dodds, R. B., and K. D. Hall. 1984. Environmental and physiological studies
of the freshwater jellyfish Craspedacusta sowerbyi. Bios 55:75-83.
42 Ted R. Angradi
Dodson, S. L, and S. D. Cooper. 1983. Trophic relationships of the freshwater
jellyfish Craspedecusta sowerbyi Lankester 1880. Limnology and
Oceanography 28:345-351.
Garman, H. 1916. The sudden appearance of great numbers of fresh-water
medusa in a Kentucky Creek. Science 44:858-860.
Howmiller, R. P., and G. M. Ludwig. 1970. A record of Craspedacusta sower-byi
in Wisconsin. Transactions of the Wisconsin Academy of Sciences, Arts
and Letters 60:181-182.
Hutchinson, G. E. 1967. A treatise on limnology: Vol. II. Introduction to lake
biology and the limnoplankton. John Wiley and Sons, Inc., New York, New
York.
Koryak, M., and P. J. Clancy. 1981. Craspedacusta sowerbyi Lankester (Hydro-zoa)
medusoid generations in two western Pennsylvania impoundments.
Proceedings of the Pennsylvania Academy of Science 55:43-44.
Koryak, M., and L. Stafford. 1981. Bubbles and jellyfish: ebullition and
Craspedacusta sowerbyi Lankester (Hydrozoa) in Tygart Lake. Unpub-lished
Report. U.S. Army Corps of Engineers, Pittsburgh District, Pitts-burgh,
Pennsylvania.
Kramp, P. L. 1950. Freshwater medusae in China. Proceedings of the Zoolog-ical
Society of London 120:165-184.
Lytle C. F. 1959. Studies on the developmental biology of Craspedacusta.
Ph.D. Thesis, Indiana University, Bloomington.
Lytle, C. F. 1962. Craspedacusta in southeastern United States. Tulane Stud-ies
in Zoology 9:309-314.
McClary, A. 1959. The effect of temperature on growth and reproduction in
Craspedacusta sowerbii. Ecology 40:158-162.
McCullough, J. D., M. F. Taylor, and J. L. Jones. 1981. The occurrence of the
freshwater- medusa Craspedacusta sowerbyi Lankester in Nacogdoches
Reservoir, Texas and associate physical-chemical conditions. Texas Jour-nal
of Science 33:17-23.
Pennak, R. W. 1989. Freshwater invertebrates of the United States. Protozoa to
mollusca. Third edition. John Wiley and Sons, New York, New York.
Reese, A. M. 1940. Craspedacusta again. American Naturalist 74:180.
Slobodkin, L. B., and P. E. Bossert. 1991. The freshwater Cnidaria — or Coe-lenterates.
Pages 125-143 in Ecology and classification of North American
freshwater invertebrates (J. H. Thorp and A. P. Covich, editors). Academic
Press, Inc., San Diego, California.
Yeager, B. L. 1987. Drainagewide occurrence of the freshwater jellyfish,
Craspedacusta sowerbyi, Lankester 1880, in the Tennessee River system.
Brimleyana 13:94-98.
Distribution and Status of Selected Fishes in North Carolina,
With a New State Record
Fred C. Rohde
North Carolina Division of Marine Fisheries
127 Cardinal Drive Extension
Wilmington, North Carolina 28405-3845
Mary L. Moser
Centerfor Marine Science Research
7205 Wrightsville Avenue
Wilmington, North Carolina 28403
and
Rudolf G. Arndt
Faculty of Natural Sciences and Mathematics
The Richard Stockton College ofNew Jersey
Pomona, New Jersey 08240
ABSTRACT - We made status surveys of 22 state-listed fishes in the
French Broad River, Nolichucky River, and Dan River systems from
1991-1995 from 105 collections at 99 sites made by us, augmented by
data from 146 collections made by others, personal communications,
and the literature. We believe state-listed endangered Polyodon spathu-la
and Percina sclera have been extirpated from the state whereas
Exoglossum maxillingua, Thoburnia hamiltoni, Noturus flavus, N.
gilberti, and Percina burtoni are either secure or rare. All six threatened
species surveyed were collected. Of these, Lampetra appendix,
Cyprinella monacha, Cottus carolinae, and Aplodinotus grunniens
probably warrant being elevated to state-endangered status, whereas
Luxilus chrysocephalus and Percina caprodes appear to be secure.
Nine species of special concern were surveyed. Noturus eleutherus and
Etheostoma simoterum are presumed to be extirpated from the state and
Acipenser fulvescens, Hiodon tergisus, and Carpiodes carpio, if not
extirpated, probably do not now have reproducing populations in North
Carolina. Etheostoma vulneratum is restricted to the Little Tennessee
River system and Percina squamata occurs in low numbers in the
French Broad, Hiawassee, Little Tennessee, and Nolichucky river sys-tems.
Both warrant consideration for elevation to threatened status.
43
44 Fred C. Rohde, Mary L. Moser and Rudolf G. Arndt
Scartomyzon ariommus and Etheostoma podostemone have apparent
healthy populations in the Dan River system. The first record of
Ichthyomyzon bdellium from North Carolina is presented.
The North Carolina Wildlife Resources Commission lists 9 species of
fishes in this state as endangered, 1 1 as threatened, and 30 as of special concern
(Article 25, Chapter 113 of General Statutes of the State of North Carolina, 1987,
amended 1991). Since 1988 we have surveyed some of these species, and to date
have reported on the distribution and status of the sandhills chub, Semotilus lum-bee,
and the pinewoods darter, Etheostoma mariae (Rohde and Arndt 1991), and
of the sharphead darter, E. acuticeps (Rohde and Arndt 1994). In this paper we
add new, and summarize existing, data on the distribution and status of 22 of the
25 state-listed fishes that occur in the North Carolina portions of the French
Broad and Nolichucky river systems (Tennessee River drainage) and in the Dan
River system (Roanoke River drainage). The former are located in the Blue
Ridge Physiographic Province and the Dan River headwaters are located in
Appalachian Mountain remnants in north central North Carolina in the Piedmont
Physiographic Province. We also provide records on a species of fish new to
North Carolina.
Three endangered, five threatened, and nine of special concern fish
species (34% of the state total) occur in the French Broad River system; two
endangered, two threatened, and one species of special concern (10% of the
total) occur in the Nolichucky River system; and three endangered and two of
special concern fishes (10% of the total) occur in the Dan River system (Table
1).
SURVEY AREAS
The French Broad River originates in North Carolina and runs some
166 river kilometers (rkm) to where it enters Tennessee and drains approximate-ly
4,163 km2 of North Carolina (Fig. 1). River elevation over this reach (Fig. 2)
drops from 640 m to 378 m; river gradient from its headwaters to Asheville is 2.6
m/km and from Asheville to the Tennessee border is 5.2 m/km (Richardson et al.
1963). Redmon Dam near Marshall in Madison County, North Carolina, pre-vents
upstream movement of fishes, and six species that might be expected to
occur farther upstream are known only from below the dam (Menhinick 1986).
The Nolichucky River and its three major tributaries (Fig. 2), the Cane,
North Toe, and South Toe rivers, drain an area of about 1,666 km2 (Crowell
1965). The Nolichucky River enters Tennessee at an elevation of 539 m (Crow-ell
1965), and joins the French Broad River at Douglas Reservoir in Jefferson
County, Tennessee (Fig. 1).
Distribution of Fishes 45
Table 1. List (1991) of endangered, threatened, and special concern fishes found
in the mountainous portions of North Carolina (from Article 25, Chapter 1 13 of
General Statutes of the State of North Carolina, 1991).
Scientific Name Common Name River System Occurrence
ENDANGERED
Polyodon spathula
Exoglossum maxillingua
Scartomyzon hamiltoni
Noturus flavus
Noturus gilberti
Percina burtoni
Percina sclera
THREATENED
Lampetra appendix
Cyprinella monacha
Hybopsis rubrifrons
Luxilus chrysocephalus
Etheostoma acuticeps
Percina caprodes
Aplodinotus grunniens
Cottus carolinae
Paddlefish
Cutlips minnow
Rustyside sucker
Stonecat
Orangefin madtom
Blotchside logperch
Dusky darter
Amer. brook lamprey
Spotfin chub
Rosyface chub
Striped shiner
Sharphead darter
Logperch
Freshwater drum
Banded sculpin
French Broad
Dan
Dan
Nolichucky
Dan
French Broad
Nolichucky
French Broad
French Broad
French Broad
Little Tennessee
Savannah
Nolichucky
Nolichucky
French Broad
New
French Broad
French Broad
SPECIAL CONCERN
Acipenserfulvescens Lake sturgeon
Hiodon tergisus Mooneye
Clinostomus funduloides Rosyside dace
Notropis lutipinnis Yellowfin shiner
Phenacobius teretulus
Carpiodes carpio
Scartomyzon ariommus
Noturus eleutherus
Etheostoma inscriptum
Etheostoma jessiae
Etheostoma podostemone
Etheostoma simoterum
Kanawha minnow
River carpsucker
Bigeye jumprock
Mountain madtom
Turquoise darter
Blueside darter
Riverweed darter
Snubnose darter
French Broad
French Broad
Little Tennessee
Little Tennessee
Savannah
New
French Broad
Dan
French Broad
Savannah
French Broad
Dan
French Broad
46 Fred C. Rohde, Mary L. Moser and Rudolf G. Arndt
Table 1. Continued.
Etheostoma vulneratum Wounded darter
Percina macrocephala
Percina oxyrhyncha
Percina squamata
Longhead darter
Sharpnose darter
Olive darter
French Broad
Little Tennessee
French Broad
New
French Broad
Hiwassee
Little Tennessee
Nolichucky
Fig. 1. Upper Tennessee River drainage, North Carolina and Tennessee.
C A R O UNA _
£.-~£o'u Na"~"
TENNESSEE /
G E O R *G I A
.- so vn H
The Dan River is the major southern tributary to the Roanoke River. It
originates on the Blue Ridge uplands in south central Virginia and, after a course
of 59 rkm, enters North Carolina in northwest Stokes County; it crosses the state
line five more times before joining the Roanoke River at Kerr Reservoir in Hal-ifax
County, Virginia (Fig. 3). The 140 rkm North Carolina portion drains
approximately 4,410 km2 of the state. River elevation drops from 366 m at the
point where it first enters North Carolina to 140 m where it first exits North Car-olina
in northeast Rockingham County (from U.S. Geological Survey 7.5 minute
series topographic maps).
Distribution of Fishes 47
Fig. 2. French Broad and Nolichucky river systems, North Carolina.
Fig. 3. Dan River system with sites (dots) we sampled from July 1992 to May
1995. Some dots overlap.
Patrick County %
TOWNESOAM ^
N
7.5 15
kilometers
PINNACLES \ POWER 4
PLANT \
JVV* TALBOTTOAM
. /LITTLE OAN RIVER T\
Homy County
VIRGINIA ~"S—
NORTH CAROUNA
Surry County
i
C DAN RIVER J
MAYO RIVER
-A ^S M MMSON \r~r-
"~^""
^^ StotosCounty V*Rockingham County
48 Fred C. Rohde, Mary L. Moser and Rudolf G. Arndt
All three rivers are generally bordered by forest, although some land is
pasture. River substrate ranges from bedrock to boulders to cobble to silt.
Fig. 4. French Broad and Nolichucky river systems with sites (dots) we sampled
from July 1991 to April 1995. Some dots overlap.
<^ p TENNESSEE / \ -^r /
Nolichucky River _•'" N^^ i "-vt L.
y "\ /**"
French Broad River -*>Jfc=-;
Cane Rfver jj.
\ North Toerfiiver
vt\
'w J
/'jm
-^
—
^-r ) L
A South Toe River
1N
) J
1 1 1
NORTH CAROLINA \f 16 32
KILOMETERS
METHODS AND MATERIALS
We sampled 99 sites in 105 collections: 32 sites in the lower reach of
the French Broad River and some of its tributaries from May 1994 to May 1995,
and at 2 sites in the middle reach of the river in September 1993 (Fig. 4); 39 sites
in the Nolichucky River system between July 1991 and November 1994 (Fig. 4);
and 15 sites in the Dan River and in 2 of its tributaries 25 times from July 1992
to May 1995 (Fig. 3). We sampled most sites only once, although four sites each
in the French Broad and Nolichucky rivers were each sampled twice. We also
sampled 1 1 sites in the Virginia portion of the upper Dan River from November
1993 to October 1994 (Fig. 3). All site locations and dates are available from the
senior author. In addition, we include data from 146 collections made by others,
other personal communications, and from the literature.
We sampled primarily with a backpack electroshocker and seine, using
the technique described by Jenkins and Burkhead (1975). Each site was elec-trofished
for 45- 190 minutes, i.e., until we believed that sampling had been com-
Distribution of Fishes 4 9
prehensive. However, in the French Broad and in the Nolichucky rivers, we sam-pled
12 sites in 1991 and 2 in 1994 only by seine (3.05 m x 1.2 m, 0.64 cm mesh),
and we sampled 2 sites in the former river in 1994 only by a 25 m, 14 cm
stretched mesh monofilament gill net and a 50 m, 5.1 cm stretched mesh
monofilament gill net. The nets were deployed overnight and fished on consec-utive
days for a total of six net days. In the Dan River in 1992 and in 1994, we
sampled two sites in each year only by seine (size as above). At five sites in the
North and South Toe rivers, in addition to sampling with electroshocker and
seine, we also surveyed fishes underwater by snorkeling. In all sampling efforts,
the known preferred habitat for each species was sampled most intensively.
In addition to the fishes taken, data on stream depth, width, and sub-strate
type; current; air and water temperatures; pH; and dissolved oxygen con-centration
were often recorded at a site, and we include these data when avail-able.
Fishes were preserved in 10% formalin upon capture for subsequent exam-ination.
Fish measurements when available are given; TL is total length and SL
is standard length. We deposited preserved specimens in the North Carolina
State Museum of Natural Sciences in Raleigh. Scientific and common names of
fishes used herein follow Mayden et al. (1992), except for Cyprinella monacha
which follows Jenkins and Burkhead (1994).
We include figures that show all of our known fish capture localities in
North Carolina. In our species accounts we occasionally include records of fish-es
taken in portions of adjacent states in an effort to make the accounts more
accurate and complete.
Positive results are gratifying, but, as usual, negative results are not nec-essarily
conclusive. When fish populations in rivers and large creeks decrease
strongly, it becomes virtually impossible to differentiate between occurrence at a
low level and extirpation (Etnier 1994).
RESULTS AND DISCUSSION
TENNESSEE RIVER DRAINAGE
ENDANGERED SPECIES
Paddlefish, Polyodon spathula (Walbaum)
The paddlefish once occurred throughout the Mississippi River and its
larger tributaries, but its distribution has decreased coincidental with river chan-nelization,
damming, and overfishing (Burr 1980). Cope (1870) maintained that
it migrated up the French Broad River as far as Asheville in Buncombe County,
North Carolina. Fishermen in this state reported that it had been caught in the
lower reaches of the French Broad River as recently as 1983, but none of these
reports has been substantiated by specimens (E. Menhinick, personal communi-cation,
1994). We sampled with large-mesh gill nets in the French Broad River
downriver of Hot Springs and in the river at the mouth of Big Laurel Creek, both
Madison County, North Carolina, on 14 and 15 May and 15 and 16 August, 1994,
50 Fred C. Rohde, Mary L. Moser and Rudolf G. Arndt
respectively. Swift currents limited our efficiency. We caught no paddlefish.
Local fishermen in Madison County whom we questioned in 1994 told us that
they had never seen a paddlefish from North Carolina nor heard of one caught
there. We presume that the paddlefish has been extirpated from North Carolina.
Fig. 5. Distribution of the stonecat, Noturus flavus, (circle) and the blotchside
logperch, Percina burtoni, (star) in the French Broad and Nolichucky river sys-tems,
North Carolina. An open circle overlaps two historical sites where the
stonecat was not taken in this survey. Specific historical sites for the blotchside
logperch in Cane Creek and the Swannanoa River are not known and are plotted
as circles with a question mark.
Stonecat, Noturus flavus Rafinesque
The stonecat is distributed through portions of the Mississippi River
basin, the Great Lakes, the Ohio River basin, and the St. Lawrence, Mohawk, and
Hudson River systems (Rohde 1980). In North Carolina it is documented from
only three sites in the Cane River, where it was collected on 1 8 June (one speci-men)
and 26 June (six), 1984, and 15 September 1985 (two) (Menhinick 1986).
Despite our efforts to collect it at all three sites, we took two adults (pho-tographed
and released) only at the downstream-most site on 4 September 1993
(Fig. 5).
Distribution of Fishes 5 -j
We record it here for the first time from the Ivy River, a tributary to the
French Broad River, from upstream of Marshall in Madison County, North Car-olina,
where we took three adults (90-99 mm SL) on 14 August 1994 (Fig. 5).
One adult (86 mm TL) and one specimen (released, not measured) were collect-ed
in the Little Tennessee River at Needmore in Swain County, North Carolina,
by Tennessee Valley Authority (TVA) personnel on 20 June 1994 (E. Scott, per-sonal
communication, 1994) (Fig. 1). This site had been sampled five times by
TVA biologists during the period 1988-1993 and once by us in 1993. The dis-covery
of the species there in 1994 was unexpected. Preferred habitat was grav-el
riffles. Current at the Ivy River site was 0.31 m/sec, water temperature 20.6
C, pH 8.0, and dissolved oxygen concentration 8.0 ppm. Its status of endangered
in North Carolina is warranted.
Blotchside logperch, Percina burtoni Fowler
The blotchside logperch occurs in disjunct populations in the Ten-nessee
River drainage from westcentral Tennessee to southwestern Virginia
(Page and Burr 1991); where it occurs it is localized and rare (Etnier 1994). In
North Carolina it was taken at one site in Cane Creek (Henderson County) in
1902, at one site in the Swannanoa River (Buncombe County) in 1934, and at
two sites in the South Toe River (Yancey County) in 1975 and 1977 (Menhinick
1986) (Fig. 5). We collected two adults (120 mm SL, one released) at a new
locality in the South Toe River near its confluence with the North Toe River in
September 1993. Since we noted that this fish can readily avoid electroshockers
and seines, we observed 2 to 6 adults in each of 4 visits by snorkeling at the two
upstream historic sites in the South Toe River in July and September 1993 and
August 1995 (Fig. 5). Preferred habitat was in pools below riffles. Menhinick
(1986) presumed P. burtoni to have been extirpated from the Swannanoa River
and from Cane Creek since he did not obtain it there in 14 collections nor did the
North Carolina Division of Environmental Management in 5 collecions (V
Schneider, personal communication, 1994). We did not collect it in either stream
in two collections made there in 1993, and we concur with Menhinick (1986). Its
continued presence in North Carolina is tenuous.
Dusky darter, Percina sciera (Swain)
The dusky darter occurs from the Wabash River drainage in Indiana
south and west to the Guadalupe River drainage in Texas and east to the Tombig-bee-
Black Warrior river system in Alabama (Page 1980). In North Carolina it is
known only from Spring Creek, Madison County, where U.S. Forest Service per-sonnel
collected one specimen in 1966 and one in 1969 (Auburn University Col-lection
3442), although the specific sites are now not known (M. Seehorn, per-sonal
communication, 1994). We made 13 collections at 7 sites with suitable
habitat in Spring Creek over a distance of 27.3 rkm in 1994 and 1995. We did
not collect it. Apparently never widespread or common in North Carolina, we
52 Fred C. Rohde, Mary L. Moser and Rudolf G. Arndt
consider the dusky darter to have been extirpated from this state by unknown
causes. It appears to be a relatively tolerant species in other portions of its range.
THREATENED SPECIES
American brook lamprey, Lampetra appendix (DeKay)
The American brook lamprey is widely distributed in the St. Lawrence
and Mississippi river basins from New York to northern Arkansas, and on
Atlantic Slope drainages from southern Quebec south to the Roanoke River
drainage in Virginia (Page and Burr 1991). In North Carolina it was known from
only one site in the downstream reach of Spring Creek, Madison County, at a
point where a railroad trestle crosses this creek, where 26 individuals were taken
in 1980 and 1 in 1983 (Menhinick 1986). We took one adult some 200 m down-stream
of the above site (and about 50 m upstream of the confluence of Spring
Creek with the French Broad River, Madison County) on 22 April 1995 (Fig. 6),
and two adults on the same day, also in Spring Creek at a point about 0.9 rkm
above this confluence. They were males and measured 144, 147, and 157 mm
TL. All were taken in gravel riffles where the current was 0.45 m/sec. We col-lected
one ammocoetes of 152 mm TL in a sandy-bottomed pool about 50 m
downstream of the last-mentioned Spring Creek site on 14 August 1994. The pH
here was 6.9, and the dissolved oxygen concentration was 8.4 ppm. This species
appears to be restricted to this creek in North Carolina. Its status of threatened
in North Carolina appears to be conservative.
Fig. 6. Distribution of the American brook lamprey, Lampetra appendix, (circle)
and the striped shiner, Luxilus chrysocephalus, (star) in the French Broad and
Nolichucky river systems, North Carolina. Some symbols overlap sites.
NORTH CAROLINA
Distribution of Fishes 5 3
Spotfin chub, Cyprinella monacha (Cope)
The spotfin chub is endemic to the Tennessee River drainage in disjunct
populations from southwestern Virginia to northwestern Alabama and in the Buf-falo
River in central Tennessee (Jenkins and Burkhead 1984, Etnier and Starnes
1994). It has apparently been extirpated from Alabama and Georgia (Etnier and
Starnes 1994). In North Carolina it is restricted to a 16.8 rkm section of the Lit-tle
Tennessee River in Macon and Swain counties (Alderman 1987). We col-lected
and released 27 spotfin chub in this river at Needmore in Swain County
(Fig. 1) on 25 September 1993. Three previous North Carolina records, two
from the French Broad River system (1888) and one from the Tuckaseegee River
(1940), apparently represent now extirpated populations (Menhinick 1986). We
did not take it in 36 collections in the French Broad River system, and we con-cur
with Menhinick (1986) that it has been extirpated from this drainage. Its sta-tus
of threatened in North Carolina appears to be conservative.
Striped shiner, Luxilus chrysocephalus Rafinesque
The striped shiner is common in the southern Great Lakes basin from
western New York and southeastern Wisconsin south through much of the Mis-sissippi
River basin almost to the Gulf of Mexico (Page and Burr 1991). A dis-junct
population was discovered in the Cane River, Yancey County, North Car-olina
in 1980 by E. Menhinick, and soon thereafter it was known from five sites
in the Cane River system (Menhinick 1986). We found it in four sites in 14.5
rkm of the middle reach of the Cane River and in two tributaries, Bald and Indi-an
creeks (Fig. 6), in 1994. Numbers taken per our collections ranged from 2-
23. Specimens ranged from 29-107 mm TL. Tennessee Valley Authority biolo-gists
took 61 individuals in one collection in the Cane River in 1992 (E. Scott,
personal communication, 1994) (Fig. 6). Preferred habitat was pools and runs.
Current ranged from 0.39-0.57 m/sec, pH 7.0-7.5, and dissolved oxygen concen-tration
6.8-8.4 ppm. Its status of threatened in North Carolina is warranted.
Banded sculpin, Cottus carolinae (Gill)
The banded sculpin inhabits mountainous areas of the Mississippi River
basin from West Virginia west to Kansas and from the Ozark Mountains south-east
to southern Alabama (Page and Burr 1991). In North Carolina it was report-ed
only from Big Laurel and Spring creeks, Madison County (Robins 1954).
Menhinick (1986) later reported it as restricted to the main stream of the French
Broad River in North Carolina near the Tennessee line and absent from the two
creeks. E. Menhinick (personal communication, 1994) took two adults and eight
juveniles with rotenone in the downstream-most 100 m of Shut-in Creek, Madi-son
County, North Carolina in July 1994 (Fig. 7). We did not collect it at two
upstream-sites in this creek in 1994. However, we did take 58 specimens in two
collections made on 14 May and 19 July 1994 throughout the lower 300 m of
54 Fred C. Rohde, Mary L. Moser and Rudolf G. Arndt
Paint Creek, Greene County, Tennessee. This creek enters the French Broad
River some 120 m downstream of the North Carolina/Tennessee line (Fig. 7). Its
status as threatened appears to be conservative.
Fig. 7. Distribution of the banded sculpin, Cottus carolinae, (circle) and the
freshwater drum, Aplodinotus grunniens, (star) in the French Broad River sys-tem,
North Carolina. A circle with a question mark indicates an undefined his-torical
site of the banded sculpin. Some symbols overlap sites.
Logperch, Percina caprodes (Rafinesque)
The logperch occurs from central Canada and the upper Mississippi
River and adjacent drainages south to the Gulf of Mexico, and on Atlantic Slope
drainages from the Hudson River south to portions of the Chesapeake Bay
drainage (Rohde et al. 1994). In North Carolina it is known from four sites in
the French Broad River between Redmon Dam and the Tennessee state line
(Harned 1979) (Fig. 8), four specimens were collected below Redmon Dam in
1986 and 1987 (Birchfield et al. 1987) (Fig. 8), and from one site in the New
River, Allegheny County (Menhinick 1986). We collected eight adults (88-132
mm SL) at four sites in the downstream reaches of the French Broad River and
at two sites in the downstream portion of Spring Creek, Madison County on 13
May; 19, 22 July; and 5 November 1994 (Fig. 8). One specimen was taken in
Distribution of Fishes 55
Fig. 8. Distribution of the logperch, Percina caprodes, (circle) and the olive
darter, Percina squamata, (star) in the French Broad and Nolichucky river sys-tems,
North Carolina. Some symbols overlap sites.
cd^ ^W^^ TENNESSEE / /'
N—I
WQJj/^hi ir*ir-iy Diuar
\^£.s-~
S -\ f**
French Broad River -*~^~,~}\
Cane River j
1 North Toeffiiver
vV*w \
r"J\
-*/—^-r ) F l/_
) r" \
/* South Toe River
1N
1 J
NORTH CAROLINA \f 1 6 32
KILOMETERS
Spring Creek 7.1 rkm upstream from its mouth in July 1994 (S. Bryan, personal
communication, 1994) (Fig. 8). Preferred habitat in the river was runs with large
boulders. Current at our Spring Creek site was 0.58 m/sec, pH 7.1-7.6, and dis-solved
oxygen concentration 7.1-10.5 ppm. Its status of threatened is warranted.
Freshwater drum, Aplodinotus grunniens Rafinesque
The freshwater drum occurs throughout the Mississippi River basin
from southern Canada and the Great Lakes to western Texas and western Flori-da
(Rohde et al. 1994). Prior to this survey, it was known in North Carolina from
six sites in the lower reaches of the French Broad River downstream of Redmon
Dam, Madison County (Harned 1979) (Fig. 7). We collected one large specimen
(305 mm TL) in a pool in Spring Creek, at a point 1 rkm upstream of its conflu-ence
with the French Broad River, on 22 July 1994, and E. Menhinick (personal
communication, 1994) took one specimen in the same month in Spring Creek at
this confluence (Fig. 7). Its status of threatened is warranted due to the lack of
juveniles in collections.
5 6 Fred C. Rohde, Mary L. Moser and Rudolf G. Arndt
SPECIES OF SPECIAL CONCERN
Lake sturgeon, Acipenserfulvescens Rafinesque
The lake sturgeon is usually found over shoals in lakes and large rivers
in central Canada and Hudson Bay and St. Lawrence River drainages, and in
much of the Mississippi River drainage south to northeastern Louisiana (Page
and Burr 1991). Eight specimens, presumably of this species, were taken from
the French Broad River near Hot Springs in Madison County, North Carolina in
1945 (Brimley 1946). An occasional lake sturgeon is still reported from Douglas
Reservoir in Jefferson County, Tennessee, but these are unsubstantiated records
(Etnier and Starnes 1994). We set 2 large-mesh gill nets of 25 and 50 m total
length in the French Broad River downstream of Hot Springs in mid-May 1995
and in the river at the mouth of Big Laurel Creek in mid-August but failed to col-lect
sturgeon. Swift current limited sampling location possibilities at the former
site and reduced gear efficiency. Local North Carolina state fishery biologists
have no reported sightings (J. Borawa, personal communication, 1994). Men-hinick
(1986) considers the lake sturgeon to have been extirpated from North
Carolina, and we concur.
Mooneye, Hiodon tergisus Lesueur
The mooneye is found in central and southern Canada and in much of
the Mississippi River basin from the Great Lakes south to the Gulf of Mexico
(Page and Burr 1991). It historically occurred in the upper reaches of the French
Broad River near Bowman's Bluff, Henderson County, North Carolina in 1902
(Smith 1907), but it is now known only from Redmon Dam to the Tennessee
state line (Menhinick 1986) based on several mooneye obtained from fishermen
in the French Broad River just above the confluence with Big Laurel Creek by
Harned (1979). We did not take it in this river in our electroshocker or gill net
collections. Its status of special concern in North Carolina appears to be conser-vative.
River carpsucker, Carpiodes carpio (Rafinesque)
The river carpsucker occurs throughout the Mississippi River basin
from Montana to Pennsylvania and south to the Gulf of Mexico (Lee and Plata-nia
1980). There is one North Carolina 1947 record from the French Broad River
near Hot Springs in Madison County (Menhinick 1986). It was also captured in
the same river in Tennessee 41 rkm downstream of the North Carolina state line
in 1979 (Harned 1979), but we failed to collect it in this study. Its status of spe-cial
concern appears to be conservative.
Mountain madtom, Noturus eleutherus Jordan
The mountain madtom occurs in disjunct populations from northwest-ern
Pennsylvania south and west through the Ohio River basin to the Red and
Distribution of Fishes 57
Ouachita river drainages in Oklahoma and Arkansas (Page and Burr 1991). The
only verified North Carolina specimens are from Spring Creek, Madison Coun-ty
and were collected in 1889 (Taylor 1969). It was also collected at two sites in
the French Broad River just upstream of Douglas Reservoir, Cocke County, Ten-nessee
(32 km downriver of North Carolina) during a 1979 TVA survey (Harned
1979). We did not collect it at any of the 34 sites we surveyed in the lower reach-es
of the French Broad River system. We concur with Menhinick (1986) that it
has been extirpated from North Carolina.
Snubnose darter, Etheostoma simoterum (Cope)
The snubnose darter is abundant in the Tennessee River drainage from
southwestern Virginia to northern Alabama (Rohde et al. 1994). The only puta-tive
extant specimen from North Carolina was reported by Cope (1870) and is
now in the United States National Museum, but it is unclear from Cope's records
whether its provenance is North Carolina or Tennessee (Menhinick 1986). Men-hinick
(1986) reported two unverified records from Laurel and Spring creeks,
Madison County, North Carolina. We collected no snubnose darter, nor did Men-hinick
(1986). M. Hopey, who made 13 collections for a general survey of the
streams in this area for the Western North Carolina Alliance in 1992, did not col-lect
it (M. Kelly, personal communication, 1994). We consider the past or pre-sent
occurrence of this darter in North Carolina to be highly doubtful.
Wounded darter, Etheostoma vulneratum (Cope)
The wounded darter is restricted to the upper Tennessee River drainage
from Virginia to Georgia (Rohde et al. 1994). It is abundant in the Little Ten-nessee
River in North Carolina (F. Rohde, personal observations). Although the
type locality is Spring Creek, Madison County, North Carolina (Cope 1870),
none has been reported from the French Broad River system in North Carolina
since then, including our survey. Harned (1979) collected one specimen in the
French Broad River in Tennessee at a point approximately 23 km downstream of
the North Carolina state line. We conclude that it has been extirpated from the
French Broad River system in North Carolina. Its status of special concern in
North Carolina appears conservative.
Olive darter, Percina squamata (Gilbert and Swain)
The olive darter is confined to the Rockcastle and Big South Fork rivers
in the Cumberland River drainage in Kentucky and Tennessee and to the upper
Tennessee River drainage (Rohde et al. 1994). There are five records from the
lower reaches of the French Broad River system (three in the main river and two
in Spring Creek) in North Carolina, and four records from the Nolichucky River
system (three in Cane River and two in North Toe River); it also occurs in the
Little Tennessee and upper Hiwassee rivers in the state (Menhinick 1991). We
58 Fred C. Rohde, Mary L. Moser and Rudolf G. Arndt
did not collect it in the French Broad River system, but we collected one juve-nile
(38 mm SL) in the Cane River on 19 July 1993, and three adults (104 mm
SL, two released) in the South Toe River on 5 September 1993 (Fig. 8). Pre-ferred
habitat is around large boulders in fast riffles. Its status of special concern
in North Carolina appears to be conservative.
NEW STATE RECORD
Ohio lamprey Ichthyomyzon bdellium (Jordan)
The Ohio lamprey occurs in disjunct populations in the Ohio River
basin, where it is uncommon (Rohde and Lanteigne-Courchene 1980). Because
of its presence in nearby Tennessee, Menhinick et al. (1974) listed its occurrence
in North Carolina as probable. However, there were no records from North Car-olina
until we took one male and three females from the mouth of Spring Creek,
Madison County, on 14 May 1994 (Fig. 9). Each was adult (220-260 mm TL),
had a well-developed digestive tract, and each female was gravid. We took
another three males (243-246 mm TL), one female (244 mm TL), and two juve-niles
(not ammocoetes, nor mature adults) (152, 153 mm TL) here on 22 April
1995, as well as two adult males (239, 248 mm TL) 1 rkm further upstream in
this creek on the same day. We took three females (235-240 mm TL) in nearby
Paint Creek, Greene County, Tennessee on 14 May 1995. All our specimens
were taken over rocky riffles with a current from 0.45-0.78 m/sec.
Fig. 9. Distribution of the Ohio lamprey, Ichthyomyzon bdellium, in the French
Broad River system, North Carolina.
TENNESSEE
Nolichucky River ^ -.
Distribution of Fishes 59
DAN RIVER SYSTEM
ENDANGERED
Cutlips minnow, Exoglossum maxillingua (Lesueur)
The cutlips minnow occurs on the Atlantic slope from the St. Lawrence
River and eastern Lake Ontario drainages south to the upper Dan River in North
Carolina (Gilbert and Lee 1980). Menhinick (1986) reported it from one site on
the Dan River in Stokes County, North Carolina, within 1.6 rkm downstream of
the Virginia state line. We found it in the North Carolina portion of this river at
four sites from the Virginia line downstream to NC Route 704 and at six Virginia
sites upstream to the Pinnacles Power Plant, over a total distance of 43 rkm (Fig.
10). Numbers (39) taken in our collections ranged from 1-6 (mean 3.9), and their
length ranged from 69-133 mm SL; most specimens were adults. This species
preferred fast-flowing runs or pools, near large rocks or boulders over sand and
gravel. Current where it was collected was 0.54-0.75 m/sec; water temperature
7.7 C (November)-22 C (July); pH 6.8-7.6; and dissolved oxygen concentration
10.6-11.8 ppm (both November). The species appears to be secure in its limited
distribution in North Carolina.
Fig. 10. Distribution of the cutlips minnow, Exoglossum maxillingua, in the Dan
River system, North Carolina and Virginia.
60 Fred C. Rohde, Mary L. Moser and Rudolf G. Arndt
Rustyside sucker, Thoburnia hamiltoni Raney and Lachner
The rustyside sucker is endemic to the upper Dan River system in North
Carolina and Virginia (Jenkins and Burkhead 1994). In North Carolina it is
known only from the 1 .4 rkm downstream-most portion of the Little Dan River
in Stokes County. Here Menhinick (1986) collected four specimens at a point
some 400 m downriver of the Virginia line in 1985 (Fig. 11). We made three col-lections
in the Little Dan River, from its confluence with the Dan River upstream
to the North Carolina/Virginia line, and took one adult (144 mm SL) in a run with
gravel and rubble substrate on 21 December 1992. In Virginia we took three
adults (1 18-142 mm SL) in the Dan River at one site located 365 m, and at anoth-er
site 914 m, downriver of the Pinnacles Power Plant on 28 November

WiM^m
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N.C. DOCUMENTS
CLEAPIMGHOUSe
AUG 2 6 1998
r»if. immt 8F NORTH CAROLINA
MUltH
number 25 july 1998
EDITORIAL STAFF
Richard A. Lancia, Editor
Suzanne A. Fischer, Assistant Editor
Stephen D. Busack, Managing Editor
Brimleyana, the Zoological Journal of the North Carolina State Muse-um
of Natural Sciences, appears twice yearly in consecutively numbered issues.
Subject matter focuses on systematics, evolution, zoogeography, ecology, behav-ior,
and paleozoology in the southeastern United States. Papers stress the results
of original empirical field studies, but synthesizing reviews and papers of signif-icant
historical interest to southeastern zoology are also included. Brief com-munications
are accepted.
All manuscripts are peer reviewed by specialists in the Southeast and
elsewhere; final acceptability is determined by the Editor. Address manuscripts
and related correspondence to Editor, Brimleyana, North Carolina State Museum
of Natural Sciences, P.O. Box 29555, Raleigh, NC 27626. Information for con-tributors
appears in the inside back cover.
Address correspondence pertaining to subscriptions, back issues, and
exchanges to Brimleyana Secretary, North Carolina State Museum of Natural
Sciences, P.O. Box 29555, Raleigh, NC 27626.
In citations, please use the full name - Brimleyana.
North Carolina State Museum of Natural Sciences
Betsy Bennett, Director
North Carolina Department of Environment,
Health, and Natural Resources
James B. Hunt, Jr., Governor
Jonathon B. Howes, Secretary
CODN BRIMD 7
ISSN 0193-4406
IN MEMORIAM
Dr. Joshua Laerm, Professor at the University of Georgia and Curator of
Zoological Collections at the University of Georgia Museum of Natural History, died
28 September 1997. He was bom and raised in Pennsylvania and received an under-graduate
degree from Pennsylvania State University and graduate degrees from the
University of Illinois. He joined the faculty at the University of Georgia in 1976.
Dr. Laerm published numerous works in systematics, mammalogy, and nat-ural
history. He was particularly interested in rare and threatened or endangered mam-mals
and contributed significantly to understanding their natural history and distribu-tion
in the Southern Appalachian Mountains. His enthusiasm for science, his prolific
contributions, and his eagerness to help colleagues will be deeply missed.
Turtles (Reptilia: Testudines) Of The Ardis Local Fauna Late
Pleistocene (Rancholabrean) Of South Carolina
Curtis C Bentley and James L. Knight
South Carolina State Museum, 301 Gervais Street,
P.O. Box 100107
Columbia, South Carolina 29202-3107
ABSTRACT- The Ardis local fauna (late Pleistocene) was collected from
a group of interconnecting sediment-filled solution cavities, located in
the Giant Cement Quarry near Harleyville, Dorchester County, South
Carolina. Fossil material from the lowermost levels and the extreme
upper layer of the deposit have been radiocarbon dated at 18,940 ± 760
and 18,530 ± 725 y.b.p., respectively. These dates are considered con-temporaneous
within present resolution. Approximately ninety verte-brate
taxa were collected from the site. Fourteen were species of turtles,
including eight not previously reported from the Pleistocene of South
Carolina. Among these is the southeasternmost occurrence of Emy-doidea
blandingii. This record, in conjunction with other vertebrate fos-sils
from the site, suggests a north-south dispersal route of species along
the Atlantic Coastal Plain during interglacial-glacial transitions. Geo-graphically
isolated eastern and western populations of Emydoidea
blandingii may have existed during the maximum advance of the Lau-rentide
ice sheet. Unusually complete fossils of large box turtles recov-ered
from the site corroborate the previously suggested synonymy of the
extinct Terrapene Carolina putnami with T c. major. The fossil turtle
community of the Ardis local fauna has no modern analogue. Like the
Ardis mammals, it comprises a "disharmonious" fauna which suggests
that, during the height of the Wisconsinan glaciation, the region experi-enced
a more equable climate than that of today .
The Ardis local fauna has yielded approximately 90 species of late
Pleistocene fossil vertebrates from the Coastal Plain of South Carolina, includ-ing
a substantial mammalian (Bentley et al. 1994), avian, reptilian, amphibian,
and fish faunas currently under study. We present data on the Ardis turtle col-lection
(Appendix I), the largest Rancholabrean fauna reported from the state.
Dobie and Jackson (1979) and Roth and Laerm (1980) reported fossils of late
Pleistocene age from Edisto Island, the only other Pleistocene fossil turtle fauna
from South Carolina described to date. Among the Edisto Island fauna were ten
Curtis C. Bentley and James L. Knight
taxa of turtles, including Gopherus sp. and Malaclemys terrapin which were not
recovered from the Ardis local fauna and possibly three species of Pseudemys, P.
floridana and/or P. concinna, and P nelsoni.
The Ardis local fauna was discovered in a large open-pit mine, operat-ed
by the Giant Cement Company, located 5 km NNE of Harleyville, Dorchester
County, South Carolina (33° 14'N, 80° 26'W). Quarry operations exposed San-tee
Limestone (middle Eocene) and the clay-rich Harleyville Formation (late
Eocene) which underlie Plio-Pleistocene surficial deposits (Ward et al. 1979,
Harris and Zullo 1991). Locally, groundwater differentially dissolved the Santee
Limestone in its upper portions, so that many solution cavities contacted and
penetrated the overlying Harleyville Formation and thereby opened several of
the cavities to the Pleistocene surface (Bentley et al, 1994). The radiocarbon
dates of the Ardis material place the time of deposition at or near the height of
the Wisconsinan glaciation (Bowen 1988, Tushingham and Peltier 1993). Fur-ther
discussion on the geology, dating methods of the Ardis fossil material, pre-vious
fossil collections from the quarry, fossil collection procedures, and a local-ity
map are available in prior publications (Bentley and Knight 1993, Bentley et
al. 1994).
TAPHONOMY
At least part of the fossil assemblage collected from inside the solution
cavities at the Ardis site appears to represent an obrution deposit, the very rapid
burial of intact organisms (Brett 1990) in which many of the specimens exhibit
incipient decay. Surface openings leading to the cavities varied from a gentle
downward slope to a vertical shaft, generally allowing the Pleistocene fauna ease
of ingress and egress. This permitted animals to enter the cavities in three dif-ferent
ways: (1) "walk-in" taxa, which may have used the site for
estivation/hibernation or as denning sites and hunting grounds, for example
muskrats, mink, and woodrats (Bentley et al. 1994); (2) "wash-in" taxa from the
surface, either alive or dead, which applies most readily to large animals known
only from isolated remains e.g., Mammut sp., Bison sp., Equus sp. (Bentley et al.
1994), that would have been unable to enter the cavities during life; and (3) "fall-in"
taxa which fell into exposed verticle shafts, fossil accumulations resulting
from this type of natural trap are well documented (e.g., Webb 1974).
Because of the interconnecting "tunnel-like" nature of the cavities, a
single episodic event could produce differing water velocities within the cavities
and different rates of deposition. Seasonal flooding, depending on the intensity,
may have simultaneously smothered living animals within the cavities and
buried or reworked those that had died just prior to, or in a preceding, deposi-tional
event. Consequently, specimens incompletely or shallowly buried during
an event with a low sedimentation rate (low energy) could be completely or par-tially
exhumed and reburied by a succeeding event. This resulted in the preser-
Turtles
vation of specimens in various orientations to the bedding planes (Fig. 1), vari-ous
degrees of disarticulation, and the occasional mixing of individual elements.
An articulated Emydoidea blandingii specimen preserved in its life position with
axial skeletal elements inside, indicates little or no decay prior to its final burial
(Fig. 2). This preservation suggests the turtle was buried quickly in a high ener-gy,
high sedimentation environment (Brett and Speyer 1990), resulting in burial
deep enough to avoid reworking during subsequent episodic events. Several
articulated turtles were collected with limbs and skulls preserved within the
shells in various orientations to the bedding plane. A high energy hydrological
environment before or shortly after death would likely explain the various orien-tations
observed in well preserved specimens. Retention and preservation of
limb elements, cervical and caudal vertebrae, and skulls inside the shell may
reflect a withdrawal by the turtles in response to a catastrophic event.
Fig. 1 Emydoidea blandingii, only the carapace (.547) was preserved, ventral
side up (side view), among clay clasts from the surrounding Harleyville Forma-tion.
This illustrates the hydrodynamic effect upon some specimens prior to final
burial.
Curtis C. Bentley and James L. Knight
Fig. 2 Complete E. blandingii (.546) in situ, with axial skeleton preserved inside
the shell, indicating a "withdrawal response" and final burial prior to any signif-icant
decay.
MATERIALS AND METHODS
Morphological terminology used in this paper is taken from Carr
(1952), Ernst and Barbour (1989), Holman (1967, 1977, 1985), Holman and
Grady (1987), and Preston (1979). Taxonomy follows Conant and Collins
(1991).
Morphological comparisons of Recent skeletons to fossil material were
made against available specimens in the Florida Museum of Natural History and
the South Carolina State Museum collections. Additionally, specimens from The
University of Michigan Museum of Zoology of Emydoidea blandingii, UMMZ
155047-155054, Clemmys guttata, UMMZ 51235, 51236, 51240-51242,
159219, 155001, 155002, and Clemmys muhlenbergii, UMMZ 77140 and,
130840 were studied.
Most of the specimens in the South Carolina State Museum collections
are deposited under the base number of S.C. 94.10. and for brevity, are refer-enced
in the text only by the digits following this base number. Specimens
accessioned separately are designated by the institutional prefix of SCSM. Fos-sil
specimens deposited in the National Museum of Natural History and the
Florida Museum of Natural History are designated by USNM and UF, respec-tively.
Turtles
SYSTEMATIC PALEONTOLOGY
Testudines
Kinosternidae
Material: 1 right xiphiplastron (.25); 16 peripherals (.26-.44); 2 humeri (.45-. 46);
2 partial jaw rami (.755-. 756).
Remarks: These fossil elements could only be identified with confidence to fam-ily.
Kinosternon subrubrum - Eastern Mud turtle (Lacepede, 1788)
Material: 3 nuchals (.11 -.13); 2 right, 3 left hyoplastra (.6-. 10); 5 left hypoplas-ia
(.1-.5).
Characters used for identification: The hyoplastron of Kinosternon subrubrum
can be separated from other North American Kinosternon and Sternotherus
because the axillary notch is narrower, and from Kinosternon baurii because the
axillary notch is wider and shallower (Holman 1985). K. subrubrum hyo- and
hypoplasia differ from Sternotherus odoratus in that the elements are shorter lat-erally
than medially in S. odoratus (Preston 1979). Characters used to identify
nuchal material are discussed by Holman (1975). In addition, nuchals of K. sub-rubrum
can be distinguished from nuchals of S. odoratus because the anterior lip
of the nuchal, viewed anteriorly, is nearly straight in K. subrubrum. Nuchals of
S. odoratus, viewed anteriorly, have a decided arc.
Remarks: The eastern mud turtle inhabits a variety of shallow slow to non-mov-ing
bodies of water with a soft substrate, such as swamps, ponds, marshes, wet
meadows, and lagoons (Ernst and Barbour 1989). Kinosternon subrubrum today
ranges from southern Massachusetts and Pennsylvania along the Atlantic coast,
to the tip of Florida and west into Texas and Oklahoma (Conant and Collins
1991). K. subrubrum is common in the area of the Ardis site today and may be
sympatric with K. baurii (Lamb and Lovich 1990).
This is the first report of this species from the fossil record of South
Carolina. Dobie and Jackson (1979) and Roth and Laerm (1980) both reported
the same single pygal bone from Edisto Island as "Kinosternon sp."
Sternotherus odoratus - Common Musk turtle (Latreille, in Sonnini and
Latreille, 1802)
8 Curtis C. Bentley and James L. Knight
Material: 3 nuchals (.22-.24); 2 right hyoplastra (.14-. 15); 1 right, 5 left
hypoplasia (.16-. 21).
Characters used for identification: Identification is based on characters provid-ed
in discussion for Kinosternon subrubrum. The hyo-hypoplastron of S. minor
can be separated from the same elements in S. odoratus because the area that
forms the bridge between the plastron and the carapace is dorsally compressed
or flattened in S. minor, and not raised as in S. odoratus. The nuchals compare
most favorably to S. odoratus, following the characters used above, and addi-tionally
exhibit a strong dorsal medial keel that is generally lacking in K. sub-rubrum.
Remarks: The common musk turtle inhabits areas very similar to that of K. sub-rubrum,
preferring slow to non-moving bodies of water with a soft bottom. It
has also been collected from fast moving, gravel bottomed, streams (Ernst and
Barbour 1989). S. odoratus occurs from southern Maine and Canada southward
through Florida and as far west as Kansas and central Texas (Conant and Collins
1991), and occurs in the area of the Ardis site today.
Chelydridae
Chelydra serpentina - Snapping turtle (Linnaeus, 1758)
Material: 2 right parietals (.50, .53); 1 left postorbital (.51); 1 right prefrontal
(.52); 1 left quadratojugal (.54); 1 partial left mandible (.55); 2 right mandibles
C56-.57); 6 cervical vertebrae (.102-. 107); 1 humerus (.63); 2 radii (.69-.70); 1
right scapulo-acromial process (.64); 1 right partial acromial process (.65); 5
femora (.58-.62); 3 ilia (.66-.68); 1 caudal vertebra (.108); 1 nuchal (.95); 1 right
1st peripheral (.71); 16 unassigned peripherals (.72-.83)(2 USNM)(2 UF); 1 left
1st costal (.84); 24 partial costals (.85-.92)(8 USNM)(8 UF); 2 associated costals
C93-.94); 2 neurals (.96-.97); 2 epiplastra (.100-. 101); 2 hypoplasia (.98-.99).
Characters used for identification: Chelydra shell material is very distinctive and
easily separated from other turtles including Macroclemys. Preston (1979) pro-vides
characters that allow the identification of fragmentary material. All listed
fossil elements compare favorably to Recent skeletal materials.
Axial and appendicular skeleton - The large size and diagnostic orna-mentation
of the Chelydra skull roof elements distinguish them from all other
turtles. Macroclemys lacks the rugose cranial ornamentation of Chelydra. A
pectoral girdle was assigned to this species based on the 90° angle between the
scapula and acromial process and on the heavily striated distal ends (Holman
1966). Femora and humeri could not be separated from Recent material of C.
serpentina, and are more robust than other genera of fresh water turtles (Holman
1964) except Macroclemys, which is generally considerably larger.
Turtles
Remarks: Snapping turtles are generally found in freshwater to brackish water
habitats with soft muddy substrate (Ernst and Barbour 1989) from eastern Cana-da
through the United States east of the Rockies south through Mexico and into
Ecuador. C. serpentina occurs today in the Ardis area.
Dobie and Jackson (1979), first reported fossil material of C. serpenti-na
from Edisto Island with additional material reported by Roth and Laerm
(1980).
Macroclemys temminckii - Alligator snapping turtle (Gray, 1855)
Material: 1 partial right parietal (.109).
Characters used for identification: The fossil parietal is identical to Recent spec-imens
of this turtle, differing from C. serpentina in that the dorsal surface is
smooth, i.e. without any of the prominent ornamentation consistently found in C.
serpentina (Fig. 3). The parietal of M. temminckii is generally more robust and
is longer with respect to width than specimens of C. serpentina of comparable
sizes. This was the only fossil element of this species collected from the site.
Because this represents a significant range extension, assignment to this species
was made only after exhaustive comparisons to the fossil and Recent collections
at the Florida Museum of Natural History and the South Carolina State Museum
negated all other possibilities.
Remarks: This is the largest freshwater turtle in North America and possibly the
heaviest in the world (Ernst and Barbour 1989). The alligator snapping turtle
often can be found in the deep waters of lakes, ponds, rivers and bayous that con-tain
abundant aquatic vegetation and muddy bottoms. This turtle is highly aquat-ic
and ranges westward from northern Florida into Texas along the Gulf Coast
and thence northward up the Mississippi Valley into Illinois, Iowa and Kansas
(Ernst and Barbour 1989).
This is the first fossil or Recent evidence of Macroclemys from South
Carolina and the Atlantic Coastal Plain.
Emydidae
Emydinae
Chrysemys picta - Painted turtle (Schneider, 1783)
Material: An individual specimen consisting of a complete carapace (missing the
3rd right marginal) and plastron (.110); 7 cervical vertebrae (.110.1 -.110.7); 1
ulna (.110.16); 1 radius (.110.17); 4 phalanges (.110.18-.110.21); 1 ungual
(.110.22); 2 partial scapulo-acromial processes (.11 0.11 -.11 0.1 2); 2 coracoids
10 Curtis C. Bentley and James L. Knight
Fig. 3 Right parietals of Macroclemys temminckii and Chelydra serpentina. A)
Recent M. temminckii. B) Ardis fossil. C) Recent C. serpentina.
3 cm
(.1 10.13-.1 10.14); 2 dorsal vertebrae (.1 10.9-. 1 10.10); complete pelvic girdle
(.110.15); 1 caudal vertebra (.110.8). An individual specimen consisting of a
nearly complete carapace and plastron (.111). An individual specimen consist-ing
of a complete carapace (missing left 8-9th peripherals and 6th neural) and
plastron (.112); partial skull and mandible (.112.1); partial hyoid process
(.11 2.2); 7 cervical vertebrae (.112.20-. 11 2.26); 2 humeri (.11 2.5-. 11 2.6); 2 ulnae
(.112.7-.112.8); 2 radii (.112.9-. 112. 10); 2 scapulo-acromial processes (.112.15-
.112.16); 2 coracoids (.112.17-.112.18); 2 femora (.112.3-.112.4); 2 tibiae
(.112.11-.112.12); 2 fibulae (.112. 13-. 112. 14); 7 metapodial elements (.112.42-
.112.48); 31 phalanges (.112.49-. 112-80); 13 unguals (.112.81-.112.91); 2 sacral
ribs (.112.92-. 112.93); partial pelvic girdle (.112.19); 3 dorsal vertebrae (.112.27-
.112.29); 12 caudal vertebrae (.112.30-. 11 2.41). An individual specimen con-sisting
of a nearly complete plastron and attached 4-7th right peripherals (.113);
skull (including inner ear ossicles) and mandible (.113.1); partial hyoid appara-tus
(.113.2); 4 cervical vertebrae (.113-.9-.113.12); 2 humeri (.113.3-.113.4); 1
scapulo-acromial process (.113.7); 2 partial femora (.11 3.5-. 11 3.6); 1 tibia
(.113.8); 2 metapodial elements (. 1 1 3. 15-. 1 13. 16); 15 phalanges (.113.17-
.113.31); 5 unguals (1 13.32-. 1 13.36); 2 caudal vertebrae (. 1 13. 13-. 1 13. 14).
Specimen with partial carapace (.114); 3 cervical vertebrae (.114.3-. 114.6); 1
Turtles 11
humerus (.114.1); 1 radius (.114.2); 1 phalange (.114.7). 2 mixed specimens
both with partial fragmented carapaces and plastra (.115); 3 cervical vertebrae
(.115.5-. 115.6); 2 humeri (.115.1 -.115.2); 2 scapulo-acromial processes (.115.3-
.113.4); 2 coracoids (. 1 15.8-. 1 15.9); 2 tibiae (.115.10-.115.il); 1 ilium (.115.12);
1 phalange (.115.13). Four individual specimens with fragmented carapace and
plastron (.116-. 119). Two mixed specimens with badly fragmented carapaces
and plastra (.120).
Isolated elements: 16 nuchals (.123-.132)(3 USNM)(3 UF); (SCSM
91.170.1) partial plastron; 10 right and 2 left epiplastra (.133-. 144); 2 entoplas-tra
(.169-.170); 10 left and 3 right hyoplastra (.145-.151)(3 USNM)(3 UF); 1
right hypoplastron and xiphiplastron (.122); 5 left and 7 right hypoplasia (.152-
.157)(3 USNM)(3 UF); 6 left and 9 right xiphiplastra (.158-.168)(2 USNM)(2
UF); 8 right and 6 left 2nd costals (.227-.240); 3 right and 1 left 3rd costals (.241-
.244); 6 right and 3 left 4th costals (.245-.253); 7 right and 8 left 5th costals
C254-.268); 4 right and 3 left 6th costal (.269-.275); 4 right and 1 left 7th costal
(.276-.280); 56 peripherals (.171-.226); 1 mandible (.761); 2 right and 1 left
mandibular rami (.758-.760); 10 humeri (.289-.298); 8 femora (.281-.288).
Characters used for identification: Identification of complete shells was based
on nuchal characters (Bentley and Knight 1993), and the alignment of the verte-bral
and pleural sulci (Ernst and Barbour 1989) (Fig. 4).
Hyoplastron - The humeral sulcus does not cross dorsally over the plas-tral
scute overlap area, as in Clemmys. Terrapene and Emydoidea have hinged
plastra. Elements of comparable size can be separated from Deirochelys reticu-laria
as the dorsal scute overlap area in C. picta is wider and more sharply curved
Fig. 4 Fossil carapace of Chrysemys picta (.110) from the Ardis local fauna.
1 2 Curtis C. Bentley and James L. Knight
distally, and the articulating surface of the epiplastron and hyoplastron between
the entoplastron and the outside edge is wider in C. picta. Trachemys and
Pseudemys are larger and more robust than C picta as adults. Young specimens
of Trachemys and Pseudemys can be separated by a less pronounced or inflated
scute overlap area and signs of incomplete ossification.
Hypoplastron - The inguinal sulcus runs diagonally to the peripherals
and into the inguinal notch in Chrysemys but is parallel to the peripherals in
Clemmys. This element can be distinguished from Deirochelys as it is less elon-gate
with respect to width in C picta, and the scute overlap area is wider. It can
also be separated from Terrapene Carolina and Emydoidea by the lack of a hinge.
Adult Trachemys and Pseudemys differ in being larger and more robust than
adults of C. picta. Young specimens of Trachemys, comparable in size to adult
C. picta and completely ossified, can be separated from C. picta by a larger
bridge with respect to the hypoplastron proper and a greatly reduced or absent
epidermal attachment scar.
Entoplastron - The humero-pectoral sulcus does not cross the entoplas-tron
as in eastern species of Clemmys and in Terrapene Carolina. It can be tenta-tively
separated from Pseudemys, Trachemys, and Emydoidea by overall size, as
specimens of the preceding genera tend to exhibit incomplete ossification when
of comparable size to adult C. picta. However, size alone is not a reliable char-acter
for this element. We were unable to separate this element from that of
Deirochelys, so entoplastra are only tentatively assigned to C. picta.
Epiplastron - This element differs from other species (except Trache-mys)
in that the anterior edge exhibits a degree of serration. This element often
is serrated in specimens of Trachemys, but the size of these specimens allows
easy separation from C. picta. Young specimens of Trachemys generally lack
this serration and have a poorly developed scute overlap area in comparison to
C. picta of comparable size.
Xiphiplastron - This element can be separated from Clemmys, Ter-rapene,
and Emydoidea by the scute overlap area, which in those genera is more
pronounced than in C. picta. Further, Clemmys muhlenbergii has a posterior
edge tapered to a point, while in Clemmys insculpta the element is longer with
respect to width, with a pronounced notch where the anal sulcus wraps over the
edge, a condition minimal or lacking in C. picta. In C picta the scute overlap of
this element is wider and more pronounced on the posterior edge than on any
examined specimens of Deirochelys. This element in Trachemys is generally
more robust in adult specimens than in C. picta, and in young specimens of com-parable
sizes the scute overlap area is much less pronounced than that of C picta.
2nd Costal - This element differs from C guttata in being approxi-mately
30% wider with respect to length, and the junction between the 2nd ver-tebral
sulcus and the 1st and 2nd pleural sulcus is located generally more distal-ly
than in C. guttata. This element differs from C. muhlenbergii by having a
Turtles 1
3
greater curvature, and the above mentioned junction point forms a distinct "T"
shape in C. muhlenbergii and dips into a general "U" or "V" shape in C. picta.
The 2nd costal of C. picta differs from C. insculpta by being completely smooth
and lacking any "tortoise-like" bulges, by being more distally flared, and by hav-ing
greater curvature of the element. It can be separated from Trachemys and
Pseudemys by a lack of sculpting and smaller size, and from Deirochelys by lack
of sculpting, a more proximal rib attachment (Jackson 1978), and the proximal
tapering of the element.
3rd Costal - Differs from Clemmys, Terrapene, Trachemys, Pseudemys,
Emydoidea, and Deirochelys in that it lacks the vertebral sulcus between the 2nd
and 3rd vertebral scutes. Additionally, in Deirochelys the rib attachment is sig-nificantly
more distal than in C picta. This 3rd element can be separated from
the 5th costal in C. picta in that it lacks the posteriorly directed curvature of the
5th costal.
4th Costal - Can be separated from all other emydid turtles in that it
exhibits alignment of the sulci between the 2nd and 3rd vertebral and pleural
scutes. This sulcal alignment occurs only in the subspecies C picta picta
(Conant and Collins 1991).
5th Costal - Can be separated from Clemmys, Trachemys, Deirochelys,
Pseudemys, Emydoidea, and Terrapene by characters given for the 3rd costal.
6th Costal - Differs from Clemmys, Trachemys, Deirochelys, Pseude-mys,
Emydoidea, and Terrapene in that it lacks the sulcus of the 3rd and 4th ver-tebral
scutes.
Peripherals, femora, humeri, and mandibular rami - These elements are
tentatively assigned to this species as they compare most favorably to Recent and
fossil material of C. picta.
Remarks: The painted turtle has a wide distribution, occurring from southern
Canada south into Mexico and across the entire continental United States (Ernst
and Barbour 1989). Lakes, ponds, and streams are typical habitats of the paint-ed
turtle. Slow to non-moving, shallow aquatic environments with soft bottoms
are favored.
The completeness of many of the painted turtles recovered from the
Ardis site are strong indicators of an obrution deposit. Chrysemys picta occurs
in the Piedmont and mountains of South Carolina today but does not inhabit the
Ardis site or any other part of the Coastal Plain. This is the first fossil record
from South Carolina.
Clemmys guttata - Spotted turtle (Schneider, 1792)
Material: An individual specimen consisting of a nearly complete shell (SCSM
93.90.1) missing only the right hypoplastron and xiphiplastron, figured in Bent-
1 4 Curtis C. Bentley and James L. Knight
ley and Knight (1993), and the following associated cranial and postcranial ele-ments;
articulated skull fragment (prefrontal, frontal, postorbital, parietal, and
supraoccipital), right maxilla, both quadrates, both opisthotics, basisphenoid, and
articulated lower jaw, 1st cervical vertebra, 2 humeri, 1 ulna, 2 coracoids, 1 ischi-um,
partial pubis and ilium, 1 sacral rib, 1 fibula, 3 phalanges, and vertebrae
fragments. An individual specimen consisting of a partial carapace and plastron
missing only the left hypoplastron and xiphiplastron (.299). Isolated skull frag-ment
(parietal, supraoccipital, both maxillae, basisphenoid, 1 quadrate, 1 postor-bital,
1 partial squamosal) (.299.1). A single sub-adult individual with partial
plastron and 2 peripherals (.12 1.1 -.12 1.2).
Isolated elements: 17 nuchals (SCSM 93.90.2-.8)(5 USNM)(5 UF); 3
right and 2 left 2nd costals (.334-.338); 6 right and 5 left 3rd costals (.339-349);
6 left and 6 right 4th costals (.350-.361); 3 right and 2 left 5th costals (.362-366);
7 right and 4 left 6th costals (367-377); 1 left 7th costal (378); 34 peripherals
(300-333); 5 right and 3 left epiplastra (379-386); 3 entoplastra (.417-.419); 5
right and 13 left hyoplastra (387-399,.432)(2 USNM)(2 UF); 3 left hypoplasia
(.400-.402); 6 right and 10 left xiphiplastra (.403-.416)(l USNM)(1 UF); 1
mandible (.757); 3 humeri (.426-.428); 6 femora (.420-.425).
Characters used for identification: Identification of the two most complete spec-imens
is discussed by Bentley and Knight (1993).
Nuchals - See Bentley and Knight (1993), for characters used to distin-guish
this from other possible identifications.
Epiplastron - Differs from C. picta in that the scute overlap area is more
robust in C. guttata, and the anterior edge is not serrated as is common with C.
picta. It can be separated from C. muhlenbergii because the bulbous area where
the gular sulcus wraps onto the scute overlap is usually considerably wider medi-ally
to laterally in specimens of C. guttata, whereas the scute overlap portion that
is posterior to the bulbous area is narrower in C. muhlenbergii than C. guttata.
The length of the scute overlap posterior to the gular sulcus is longer than that of
C. picta. Also, the epidermal attachment scar in C. muhlenbergii is deeply
incised and tends to undercut the scute overlap area. The epiplastron of C. gut-tata
is rarely incised to this extent. The epiplastra of C. insculpta differ from C.
guttata in that the bulbous area where the gular sulcus crosses the dorsal surface
is only slightly or not at all bulbous in specimens of similar size. The epiplas-tron
of Deirochelys, Trachemys and Pseudemys that fall within the size range of
C. guttata have less pronounced scute overlap area compared to that of C. gutta-ta.
E. blandingii specimens of comparable size show sub-adult traits (incomplete
ossification) and have a less pronounced scute overlap area. Terrapene epiplas-tra
differ in that the area posterior to the scute overlap is concave, forming a
depression posteromedially to the gular sulcus, generally absent in C. guttata.
Turtles 1
5
Hyoplastron - Differs from Terrapene and Emydoidea in that C. gutta-ta
lacks the hinge components. C. muhlenbergii differs slightly from C. guttata
in that the scute overlap is narrower in C. muhlenbergii. Specimens of Trache-mys
and Pseudemys that fall within the size range of C. guttata have scute over-laps
that are greatly reduced in comparison to C. guttata and the elements exhib-it
incomplete ossification. Deirochelys and Chrysemys picta can be separated
from Clemmys guttata because the distance between the entoplastron and
hypoplastron is ca. 40% greater in adults of the former two genera. Also, in C.
picta, the humeral sulcus does not cross dorsally over the scute overlap area. C.
insculpta exhibits incomplete ossification when elements fall within the size
range of C. guttata.
Hypoplastron - This element can be separated from other genera of
emydid turtles by the lack of hinge components (separating it from Terrapene
and Emydoidea), or by its posterior width being greater than its length (separates
it from Trachemys and Pseudemys). Holman (1977) gives characters used to sep-arate
this element from C. muhlenbergii and C. insculpta.
Xiphiplastron - Separation of this element from C. picta is listed under
that species. It can be separated from C. muhlenbergii and C. insculpta in that the
posterior edge is generally squared off rather than tapering to a point, as in the
other two species. However, some specimens of C. guttata do exhibit a pointed
condition. These still can be separated from C. muhlenbergii because the area
where the abdominal muscle attaches to the xiphiplastron is more pronounced.
C. insculpta can also be separated from C. guttata because, in the area where the
anal sulcus crosses onto the scute overlap area, the xiphiplastron is deeply
notched, being greatly reduced or lacking in C. guttata.
Entoplastron - In C. guttata the humero-pectoral sulcus crosses the
entoplastron within the anterior half of that element. In C. muhlenbergii the
humero-pectoral sulcus may cross the entoplastron at its posterior extremity, but
typically it does not cross the entoplastron at all (Bentley and Knight 1993).
These fossil entoplastra are tentatively assigned to C. guttata, and not C. insculp-ta,
because this element is generally more robust in C. insculpta and the humero-pectoral
sulcus crosses the entoplastron more posteriorly in C. insculpta than in
C. guttata.
Costal - Characters used to differentiate costal elements from C. picta
are described in that section. Costals of Clemmys guttata differ from C. muh-lenbergii
in having substantially more curvature. The dorsal surface of costals in
C. guttata is smooth, lacking any exterior bulges or sculpturing common to adult
C. muhlenbergii, C. insculpta, Deirochelys, Trachemys, and Pseudemys. The
costals of Emydoidea, Trachemys, and Pseudemys exhibit immature traits when
they are within the size range of C. guttata. Terrapene has deeply incised sulci,
and its elements tend to be more acutely angled proximally and exhibit "bul-bous"
sculpturing.
1 6 Curtis C. Bentley and James L. Knight
Peripherals, femora, humeri, and mandible - These elements are tenta-tively
referred to this species because they compare most closely to Recent and
fossil C guttata.
Remarks: The spotted turtle ranges from northern Illinois into Ohio and Ontario,
east to Maine and New York, and south along the Atlantic Coastal Plain into
northern Florida (Conant and Collins 1991). Clemmys guttata occurs most com-monly
in bogs or marshy pastures, but it also can be found in woodland streams.
It favors habitats with soft substrates. C. guttata is frequently found away from
water, but even so it is the least terrestrial of the three eastern species of Clem-mys
(Ernst and Barbour 1989). The fossil spotted turtle remains from the Ardis
local fauna represent the oldest known material of this species (Bentley and
Knight 1993), and the first fossil record for the eastern United States. Holman
(1990) reports a right epiplastron from a 6,000-year-old fauna near Lansing,
Michigan. Interestingly, Ernst and Barbour (1989) noted a relationship between
this turtle and the burrows of muskrats, which the turtles apparently use for esti-vation
and hibernation sites. Fossil muskrats were the most common mammals
found at the Ardis site (Bentley et al. 1994) and are believed to have used the
solution cavities as burrow sites. This may help to explain the abundance of this
turtle at the Ardis site.
Clemmys muhlenbergii - Bog turtle (Schoepff, 1801)
Material: An individual consisting of a partial carapace (nuchal, 1st and 2nd left
costals and peripherals, 2 peripherals, numerous shell fragments) and plastron
(both epiplastra, and partial hyoplastron) (.429).
Isolated elements: 2 nuchals (.430-.431) ; 2 right epiplastra (.433-.434).
Characters used for identification: C. muhlenbergii fossils were distinguished
from other emydid turtles based on characters listed in previous sections, along
with additional nuchal and sulci characters given by Bentley and Knight (1993)
(Fig.5).
Remarks: The soft bottoms and slow moving waters of swamps, bogs and
marshes are typical aquatic habitats of the bog turtle (Ernst and Barbour 1989),
but this turtle can also be found on land. Clemmys muhlenbergii has a patchy
distribution in the Northeast, and ranges as far south as northern Georgia and
extreme northwestern South Carolina. This disjunct spatial pattern has been
interpreted as suggesting a larger former range (Smith 1957). The fossil evi-dence
from the Ardis site suggests that the species' range extended at least 250
km farther southward during the late Pleistocene.
Turtles 1
7
This is the second report of fossil material of C. muhlenbergii (Holman
1977) and represents the first fossil record from the eastern United States south
of Allegany County, Maryland. This is the first sympatric occurrence of C. muh-lenbergii
and C. guttata in the fossil record.
Fig. 5 Partial fossil carapace of Clemmys muhlenbergii (.429). from the Ardis
local fauna.
3 cm
1 1 1 1
Terrapene Carolina major- Gulf Coast Box turtle (Agassiz, 1857)
Material: An individual male specimen consisting of a partial carapace (SCSM
91.165.1) lacking it's anterior edge (Fig. 6), complete plastron (SCSM 91.165.2),
partial skull (SCSM 91.165.19), 8 cervical vertebrae (SCSM 91. 165. 10-. 17), 1
humerus (SCSM 91.165.5), both scapulo-acromial processes (SCSM 91.165.6-
.7), both coracoids (SCSM 91.165.8-.9), complete pelvic girdle (SCSM
91.165.3), 1 femur (SCSM 91.165.4), 1 sacral rib (SCSM 91.165.18). An indi-vidual
female specimen consisting of a complete carapace (SCSM 91.163.1) and
plastron (SCSM 91.163.2), 2 cervical vertebrae (SCSM 91.163.10-.il), both
scapulo-acromial processes (SCSM 91. 163. 4-. 5), both coracoids (SCSM
91.163.8-.9), both femora (SCSM 91.163.6-.7), and complete pelvic girdle
(SCSM 91.163.3). An individual female specimen consisitng of a complete
carapace and plastron (SCSM 9 1.1 64.1 -.2), 2 cervical vertebrae (SCSM
91.164.9-.10), 1 humerus (SCSM 91.164.8), 1 scapulo-acromial process (SCSM
91.164.4), both coracoids (SCSM 91.164.5-.6), 1 femur (SCSM 91.164.7), com-plete
pelvic girdle (SCSM 91.164.3), and 1 caudal vertebra (SCSM 91.164. 11).
An individual male specimen with complete carapace (SCSM 91. 166. 1) and pos-terior
half of plastron from hinge (SCSM 91.166.2). An individual female spec-
1 8 Curtis C. Bentley and James L. Knight
imen consisting of a partial carapace and one half of posterior plastron from the
bridge (SCSM 91.168.1). An individual female specimen consisting of a com-plete
carapace and plastron (.435-.435.1). A individual partial carapace (SCSM
91. 167.1).
Isolated elements: 8 nuchals (.459-.466); 2 right 1st costals (.471 -.472);
1 fused left 7th and 8th costal (.473); 5 costals (.474-.478); 3 left 5th peripherals
(.479-.481); 2 fused peripherals (1 USNM) (1 UF); 32 peripherals (.510-.541)(2
USNM)(2 UF); 2 complete, 10 partial anterior plastral lobes (.467-.469)(6
USNM)(3 UF); 1 right epiplastron (.470); 2 entoplastra (.508-.509); 2 complete,
9 partial posterior plastral lobes (.507)(5 USNM)(5 UF); 32 large shell fragments
(.510-.541); 1 partial skull (.436); 1 right maxilla (.437); 3 postorbitals (.438-
.440); 4 mandibles (.441-.444); 3 cervical vertebrae (.456-.458); 4 humeri (.445-
.448); 4 femora (.449-.451); 4 ilia (.452-.455).
Characters used for identification: The more complete specimens are easily sep-arated
from other emydid turtles based on their hinged plastra and overall mor-phology.
Emydoidea differs from the box turtle by its smooth, unkeeled carapace
and tends to be anteriorly constricted (Holman 1985).
Plastron - The hinged plastral elements prevent confusion with any
other emydid of North America except E. blandingii. The anterior end of this
attachment area between the carapace and plastron differs from E. blandingii, as
the carapace and plastron of Terrapene have a heavily sutured interlocking pro-trusion
and pocket respectively. In Emydoidea this area lacks the sutured "ball
and socket" mechanism and instead has a pronounced lateral flare generally
located on the 5th marginal. The large size and robust nature of the fossil ele-ments
suggest an affinity to this subspecies.
Peripherals - These elements are distinguished by an upwardly curved
anterior edge, forming, in some specimens, a "gutter-like" effect.
Humeri, femora, and ilia - Humeri and femora were separated from
other emydid turtles based on comparison to Recent specimens and characters
provided by Holman (1967, 1975). Ilia of T. Carolina have a distinctive
"boomerang-shape" and are straighter in other species (Holman 1977). Addi-tionally,
these fossil elements were identical to those retrieved from within the
shells of the more complete fossil T. c. major collected from the Ardis site.
Remarks: This is the largest of the North American box turtles and today ranges
from the coast of eastern Texas eastward along the Gulf coast into the Florida
panhandle (Carr 1952). The Gulf Coast box turtle is commonly found in marsh-es,
palmetto-pine forests, and upland hammocks. It enters water with a frequen-cy
similar to T. c. Carolina (Carr 1952, Conant and Collins 1991).
Turtles 1
9
Fig. 6 Partial fossil carapace of Terrapene Carolina major (SCSM 91.165.1)
which contained the skull (SCSM 91.165.19) shown in figure 6d.
Large fossil box turtle remains have generally been referred to as T. c. putnami
or T. c. putnami x major (Milstead 1969). T. c. putnami differs from T. c. major
only in size, attaining lengths upwards of 300 mm (Auffenberg 1967, Milstead
1969). The largest Recent specimen of T. c. major on record has a carapace
length of 216 mm (Conant and Collins 1991). Auffenberg (1967) reported a
large box turtle (233 mm), with skull, from Haile 8A, stating that it was very sim-ilar
to T. c. major. Blaney (1971) placed T. c. putnami in synomyny with T. c.
major, based on the shared characters of the two subspecies, and stated that size
alone was not a justifiable reason to recognize a subspecies. The fossil box tur-tles
from the Ardis site exhibit all the characters Milstead (1969) used to distin-guish
T. c. putnami. Furthermore, the five specimens had carapace lengths of
190.0 mm to 260.0 mm. Shell and axial elements of fossil box turtles from the
Ardis site could not be consistently distinguished from Recent specimens of T. c.
major except by size. Milstead (1969) stated that populations of T. c. Carolina in
Massachusetts and Michigan, on the northwestern edge of the subspecies range,
appear to have strong morphological affinities to T. c. major having average cara-pace
lengths of 140 mm and 139 mm respectively. Milstead suggested that this
relationship may be due to a pre-Wisconsin influence of T. c. putnami or the
influence of T. c. triunguis. An isolated, fused posterior plastral lobe (82.26 mm)
collected from the Ardis site was estimated to belong to a specimen with a cara-pace
length of about 140-145 mm. This plastral lobe might indicate the presence
20 Curtis C. Bentley and James L. Knight
of box turtles at or near the size of the northwestern populations discussed by
Milstead (1969). Additionally, several significantly smaller isolated peripherals
were collected from the Ardis site, but these peripherals are largely unfused and
may represent juveniles. The association of the smaller fused plastral lobe with
significantly larger specimens suggests that during the height of the Wisconsin
glaciation there may have been intergradation between local and northerly dis-placed
populations of T. c. Carolina, and populations of T. c. major, T. c. triun-guis,
and T. c. baud radiating from the south. The Ardis population, being pre-dominantly
large box turtles, suggests that T. c. major traits (very large size,
elongated shells, and upward curvature of peripherals) were more predominant
during this time period.
The Ardis site produced two partial fossil skulls; SCSM 91.165.19 asso-ciated
with a carapace of 245-250 mm. in length, and (.436), an isolated partial
skull comparable in size to SCSM 91.165.19 (Fig. 6 and Fig. 7). We failed to
distinguish any consistent differences between our skulls and Recent specimens
except for size and an exaggerated upward curvature of the supraoccipital crest
in specimen SCSM 91.165.19. This extreme curvature is considered an anom-aly
due to its complete absence in other fossil specimens; however, it is noted in
Recent specimens but is less developed. The supraoccipital is thought to be one
of the most variable cranial elements for box turtle systematics (W. Auffenberg,
University of Florida, personal communication).
The parietals of most Recent specimens examined of Terrapene Caroli-na
had parietals that were inflated anteriorly, with a reduction in the tabled, dor-sal
surface of this element. Although extremely preliminary, we suggest a cor-relation
between size and the degree of parietal inflation in T. c. major. Anteri-or
inflation of the parietals is greatly reduced to absent among the largest speci-mens.
In other subspecies of T. Carolina, the inflation of the parietals remains
fairly constant. The two skulls from the Ardis site do not exhibit anterior infla-tion
of the parietals. The morphological similarities between the fossil box tur-tles
of the Ardis site and the Recent and fossil specimens examined from muse-um
collections (Fig. 7) suggests that the greatest affinity of the Ardis specimens
is to T. c. major. They support the synonymy of T. c. putnami with T. c. major
(Blaney 1971). Affinities noted by Milstead (1969) between northwestern pop-ulations
and T. c. major may be a result of the proposed intergradation between
box turtles during the height of the Wisconsin glaciation. The synonymy of T. c.
major with T. c. putnami also suggests that T. c. major may have been capable of
obtaining a considerably larger size than that observed in living specimens.
Although we believe a systematic revision of the genus Terrapene is
needed, it exceeds the bounds of this faunal review. One of the goals of this dis-cussion,
however, is to emphasize the need for such a revision, based both on
"standard" characters and on osteological characters of all fossil and extant
forms.
Turtles 21
Fig. 7 Terrapene Carolina major skulls. A) Recent (UF 18963). B) Haile 8A (UF
3148). C) Ardis fossil (SCSM 91.165.19). D) Ardis fossil (.436).
3 cm
3 cm B
3 cm D
22 Curtis C. Bentley and James L. Knight
Deirochelys reticularia - Chicken Turtle (Agassiz, 1857)
Material: 2 peripherals (.542-.543).
Characters used for identification: Identification is based upon the distinctive
"spike-like" pattern of the dorsal sculpturing on these elements (Jackson 1964,
Holman 1978).
Remarks: This turtle today has a continuous range from Texas and Oklahoma
through the Gulf states and along the southern half of the Atlantic Coastal Plain
states into North Carolina, with isolated populations in southeastern Virginia
(Conant and Collins 1991). Still-water habitats, such as ponds, swamps, marsh-es,
and temporary pools, are commonly occupied by chicken turtles, which
reportedly do not favor moving water (Ernst and Barbour 1989).
This is the first fossil record of Deirochelys from South Carolina. The
species most extensive fossil record and probable origin is in Florida (Jackson 1978).
Emydoidea blandingii - Blanding's Turtle (Holbrook, 1838)
Material: An individual specimen consisting of a complete carapace, anterior
plastral lobe (.544), left lower jaw (.544.1), partial hyoid (.544.2), 1 partial
humerus (.544.4), 1 partial scapulo-acromial process (.544.3), 1 femur (.544.5),
3 partial dorsal vertebrae (.544.8-.544. 10), 1 ilium (.544.6), 1 pubo-pectineal
process (.544.7), 1 sacral rib (.544.11). An individual specimen consisting of a
complete carapace, 1 phalange, 2 partial vertebrae (UF). An individual specimen
consisting of a nearly complete carapace, anterior plastral lobe, 4 partial dorsal
vertebrae, 3 caudal vertebrae (USNM). An individual specimen consisting of a
complete carapace, posterior plastral lobe (.545), 5 cervical vertebrae (.545.1-
.545.5), 2 humeri (.545.21-.545.22), 1 ulna (.545.31), 2 scapulo-acromial
processes (.545.27- .545.28), 2 coracoids (.545.25-.545.26), 1 dorsal vertebra
(.545.6), 2 femora (.545.23-.545.24), 1 fibula (.545.29), 1 tibia (.545.30), 1 com-plete
pelvic girdle (.545.20), 5 phalanges (.545.32-.545.35), 1 ungual (.545.36),
2 sacral ribs (.545.37-.545. 38), 13 caudal vertebrae (.545.7-.545. 19). An indi-vidual
specimen consisting of a complete carapace and plastron, partial skull (2
maxillae, 1 quadrate, basioccipital-condyle, basisphenoid, frontal-postorbital-parietal
skull fragment) (.546), partial hyoid apparatus (.546.2), 6 cervical verte-brae
C546.3-.546.8), 1 humerus (.546.20); 2 ulnae (.546.25-.546.26), 1 radius
(.546.27), 2 scapulo-acromial processes (.546.28-.546.29), 2 coracoids (546.30-
.546.31), 1 femur (.546.21), 1 tibia (.546.22), 2 fibulae (.546.23-.546.24), com-plete
pelvic girdle (.546.19), 10 phalanges (.546.32-.546.41), 1 ungual (.546.42),
2 sacral ribs (.546.43), 10 caudal vertebrae (.546.9-.546. 18). An individual spec-imen
consisting of a complete carapace (.547). An individual specimen with the
Turtles 2 3
anterior one half of the carapace (.548). An individual specimen with the ante-rior
two-thirds of the carapace (.549). An individual juvenile specimen consist-ing
of a partial carapace (nuchal, 1-2,4,8 neurals, pygal, 2-5, 7-11 left peripher-als,
1-4 right peripherals, 3-8 right costals, 3rd left costal) and anterior lobe of
plastron missing left epiplastron (.550), left xiphiplastron (.550.1), 1 dorsal ver-tebra
(.550.2).
Isolated elements: 1 nuchal (.588); associated nuchal and 1st left and
right costal with 1st left peripheral (USNM), 4 associated costals and single
peripheral (.566); 1 left and 1 right 1st costal (.567-.568); 1 left 3rd costal (with
rodent gnaw marks) (.569); 10 costals (.556-.565); 6 neurals (.570-.575); 3 right
1st peripherals (.576-.578); 1 left 5th peripheral (sub-adult) (.580); 1 left 6th
peripheral (.579); 1 partial plastron (.607); 2 anterior plastral lobes (2 UF), 1 par-tial
anterior plastral lobe (USNM), 2 pairs of associated epiplastra (.589-. 590); 4
left and 2 right epiplastra (3 USNM)(3 UF), 7 entoplastra (.581-.587); 4 left and
6 right hyoplastra (.601-.602)(4 USNM)(4 UF); 4 posterior plastral lobes
(.591)(1 USNM)(2 UF); 1 associated right hypoplastron and xiphiplastron
(.592); 7 right and 8 left hypoplastra (.593-.600)(3 USNM)(4 UF); 3 right and 3
left xiphiplastra (.603-.606)(l USNM)(1 UF); 2 partial skulls (.551-.552); 1 left
postorbital (.553); 2 left auditory bullae and quadrate (.554-.555); 5 cervical ver-tebrae
(.614-.618); 3 humeri (.608-.610); 3 femora (.611-.613); 1 partial pelvic
girdle (.619).
Characters used for identification: It is possible to distinguish the more complete
specimens of Emydoidea blandingii from Deirochelys reticularia on the basis of
the hinged plastral elements and the lack of carapacial sculpturing in the former
(Jackson 1978). Characters that distinguish this species from Terrapene Carolina
are given under that account. Isolated specimens can be distinguished from other
emydid turtles based on characters mentioned in other sections of our paper and
the following:
Epiplastron - Emydoidea epiplastron can be contrasted to Terrapene
epiplastron in several ways. This element in Emydoidea differs from Terrapene
by the presence of a depression located on the dorsal surface and medially to the
anterior scute overlap area, which is less pronounced in Emydoidea. When com-pared
to specimens of Terrapene of comparable size, this element is less robust
and somewhat dorso-ventrally compressed. However, there is some difficulty in
distinguishing large specimens of T Carolina major from E. blandingii. This ele-ment
differs from Trachemys in that it is more elongated and thinner in E.
blandingii.
Xiphiplastron - This element is most easily confused with Clemmys.
Preston and McCoy (1971), suggested that the xiphiplastron of Clemmys is wider
with respect to its length. Preston (1979) discusses additional characters used to
identify this element in Emydoidea.
24 Curtis C. Bentley and James L. Knight
Neurals - These thin, very broad, smooth elements are distinctive
among emydid turtles.
Axial and appendicular skeleton - Although cranial material of this
species is distinctive, the generalized nature of the postcranial material makes the
assignment to E. blandingii tentative.
Remarks: The recent distribution of E. blandingii is limited to southern Ontario
and the Great Lakes region, with scattered populations occuring westward into
northeastern Nebraska and south into northeastern Missouri, and eastward into
New York and Massachusetts on the Atlantic coast (Conant and Collins 1991).
The nearest fossil records of Emydoidea blandingii to the Ardis site are from
Catalpa Creek, Mississippi (Jackson and Kaye 1975), and at New Trout Cave,
West Virginia (Holman and Grady 1987). Both records are late Pleistocene.
The well preserved fossil material from the Ardis site (Fig. 8) is the first
report of this species on the Atlantic Coastal Plain and is a range extension of
about 1,200 km from its present continuous distribution and nearly 525 km south
of the nearest reported fossil locality at New Trout Cave.
Habitats frequented by E. blandingii are generally in shallow, lentic
waters with soft substrate, such as ponds, streams, marshes and sloughs (Ernst
and Barbour 1989).
Fig. 8 Fossil Emydoidea blandingii carapace (.547).
5 cm
Turtles 2 5
Trachemys scripta - Slider Turtle (Schoepf, 1792)
Material: SCSM 91.169.1 a nearly complete carapace and plastron, partial right
lower jaw, 1 scapulo-acromial process, 1 coracoid, 1 partial vertebra, partial
pelvic girdle, 1 fibula. An individual specimen consisting of a partial carapace
and plastron (.620).
Isolated elements; 3 nuchals (.621 -.623); 23 partial costals (.624-
.640)(3 USNM)(3 UF); 15 peripherals (.641-.655); 1 suprapygal (.656); 10 neu-rals
C657-.666).
Characters used for identification: This turtle is distinguished from other turtles
by characters given by Holman (1985). Additionally, it has a diagnostic carapa-cial
ornamentation.
Remarks: The slider has a nearly continuous distribution from Illinois southward
into Texas and New Mexico and thence eastward across northern Florida north
along the Atlantic coast into Virginia, with populations in Mexico and Maryland
(Conant and Collins 1991). Trachemys scripta can be found in most freshwater
habitats, but seems to prefer slow to non-moving water with a soft substrate
(Ernst and Barbour 1989).
Today Trachemys scripta is common around the Ardis site and we have
observed more than 10 within 100 m of the excavation site.
Pseudemys sp.-Cooters (Gray, 1855)
Material: 1 right 1st peripheral (.667); associated 9- 10th right peripherals (.668);
2 peripherals (.669-.670).
Characters used for identification: These compare most favorably to species in
this genus based on the thin, elongated sloping nature of the elements. We were
unable to identify any diagnostic characters on these elements that could be used
make a species placement. The only genus with which these may be easily con-fused
is Trachemys. The fossil elements lack the sculpturing found in Trache-mys
and are significantly thinner and more elongated. The fossil peripherals also
have a straight distal margin which contrasts with the notched margin in Trache-mys.
26 Curtis C. Bentley and James L. Knight
Remarks: Pseudemys is a common genus of the Southeast, found in various
aquatic systems (Conant and Collins 1991). The two species found in South Car-olina
today are P. floridana and P. concinna.
Testudinidae
Hesperotestudo crassiscutata -Giant Tortoise (Williams, 1950)
Material: 9 carapacial and plastral fragments (.671-.677)(1 USNM)(1 UF); 1
right xiphiplastron (.678); 1 vertebra (.685); 1 ungual (.686); 6 osteoderms (.679-
.684).
Characters used for identification: The fragmented shell elements were assigned
to this species rather than H. incisa on the basis of their extremely large size and
robustness (40.0 mm thick). The large osteoderms, phalange, and vertebra could
not be distinguished from H. crassiscutata in the Florida Museum of Natural
History.
Remarks: Bramble (1971), Preston (1979), and Meylan (1995) place all North
American non-Gopherus tortoises into the genus Hesterotestudo, and that prac-tice
is followed here. Dobie and Jackson (1979), provided the first report of
(Geochelone) H. crassiscutata in South Carolina from the late Pleistocene of
Edisto Island.
Trionychidae
Apalone sp.- Softshell turtle (Rafinesque, 1832)
Material: 1 partial nuchal (.47); 1 costal distal end (.48); 1 partial neural (.49).
Characters used for identification: These fossils are easily assigned to this genus,
based on the relatively thin shell elements with characteristic pitting of the dor-sal
surfaces, general morphology, and the geographical distribution of Triony-chidae.
However, based on these elements, we were unable to identify a species
with any certainty.
Remarks: Two species of the genus Apalone now occur in South Carolina, A.
ferox and A. spinifera. Both species inhabit various aquatic environments with
muddy or sandy bottoms in deep or shallow water (Ernst and Barbour 1989).
Apalone ferox occurs throughout Florida and in the southern portions of Alaba-ma,
Georgia, and South Carolina. A. spinifera is restricted in Florida to rivers in
the extreme northeastern and northwestern portions, and ranges no farther north
along the Atlantic coast than North Carolina. It ranges westward into Colorado
and north into Minnesota, with isolated populations in Montana, California, and
Turtles 2 7
Mexico (Conant and Collins 1991). Only A. spinifera inhabits the area of the
Ardis site today. Dobie and Jackson (1979) reported "Trionyx sp." from Edisto
Island, South Carolina. Meylan (1987) has shown the correct name for North
American softshells to be Apalone.
DISCUSSION AND PALEOECOLOGY
The turtle assemblage of the Ardis local fauna provides important new
data for the late Pleistocene of the southeastern United States. In particular, it
documents a shift in the spatial patterns of several turtle species during the late
Pleistocene. The turtle fauna is unique in its geographical and temporal setting,
and it contains the first sympatric fossil occurrences of several taxa, e.g., Clem-mys
muhlenbergii, C. guttata, Emydoidea blandingii, Macroclemys temminckii.
All of the fossil turtles collected from the site, except Hesperotestudo
crassiscutata and Terrapene Carolina major, are primarily aquatic and are com-monly
found in, or require, still or slow moving water with a soft substrate and
aquatic vegetation (Ernst and Barbour 1989). Additional evidence of a nearby
body of water included the presence of Alligator mississippiensis and elements
of the fish fauna which are currently under study. This agrees with the habitat
suggested by Bentley et al. (1994) based on the Ardis mammal fauna that indi-cates
an ecotone between a mixed forest of conifers, hardwoods and meadows,
and a permanent body of water such as a river or stream which may have given
way to a bog or marsh. Portions of the mammalian fauna and avian material
from the Ardis site further suggest the presence of a nearby large body of water
such as a lake or pond. This association is based on the life histories and habitat
requirements of many of the extant species represented in the Ardis fauna.
The Ardis turtle fauna consists of thirteen extant and one extinct
species, with five taxa considered extralimital. Three of the five have northern
affinities: Emydoidea blandingii, Clemmys muhlenbergii, and Chrysemys picta.
Terrapene Carolina major has a strong southern affinity. Macroclemys has a pri-marily
Gulf coast distribution but extends as far north up the Mississippi Valley
as Iowa (Pritchard 1989). Although Hesperotestudo was widespread in North
America by the Miocene, Hibbard (1960) suggested that the presence of Hes-perotestudo
crassiscutata in a fauna indicated a mild climate with frost-free win-ters.
The sympatric occurrence of species that are apparently ecologically
incompatible today constitutes a "disharmonious fauna" (sensu Lundelius et al.
1983), which has been interpreted by many authors (Hibbard 1960, Holman
1980, Lindelius et al. 1983) as indicating a more equable climate (reduced sea-sonal
temperature gradients) than that experienced in the region today. The
Ardis local fauna reflects such a disharmonious biota, which clearly has no mod-ern
analogue. The turtle fauna corroborates conclusions made on the basis of the
Ardis mammal fauna (Bentley et al. 1994), which also suggests that a more
28 Curtis C. Bentley and James L. Knight
equable climate than today prevailed near or during the maximum advance of the
Laurentide ice sheet in the southeastern United States.
Based on the distribution of terrestrial vertebrates, Smith (1957) stated
that the northeastern biota of North America was displaced southward during the
Wisconsinan glacial maximum. Smith also suggested that during a "Climatic
Optimum," southern counterparts dispersed northward. During the height of the
Wisconsinan glaciation, E. blandingii would have been extirpated from its pre-sent-
day northerly distribution (Mickelson et al. 1983, Conant and Collins
1991). The southern boundary of the Laurentide ice sheet, while abutted against
the Appalachian Plateau (Mickelson et al. 1983), could potentially have forced
the range of E. blandingii to be split, with one population occurring along the
Mississippi Valley and another along the Atlantic Coastal Plain. This may have
produced geographically isolated eastern and western populations of Emydoidea
blandingii during the late Pleistocene.
In addition to Emydoidea blandingii, Spermophilus tridecemlineatus
(thirteen-lined ground squirrel) was also collected from the Ardis site (Bentley et
al. 1994). Based on Recent and fossil distributions (Kurten and Anderson 1980),
the present northeastern distribution of the thirteen-lined ground squirrel also
may have been displaced southward along the Atlantic Coastal Plain by the
advancing Laurentide ice sheet.
As with the Gulf Coast Corridor, depressed sea levels during the Pleis-tocene
glacial stage exposed much or all of the Atlantic continental shelf (Bloom
1983), thereby widening the Atlantic Coastal Plain. This may have facilitated
the dispersal of glacially-displaced species southward along the coast. The
newly emergent land area provided expanded habitats for species to utilize
(Blaney 1971), and a more equable climate may have allowed for the range
extention of both northern and southern species into these new areas. Northern
populations of T. c. Carolina in Massachusetts and Michigan that show affinity to
T. c. major, discussed by Milstead (1969), may be relicts left over from a Pleis-tocene
interval of extensive intergradation between the northerly displaced sub-species
and radiating southern subspecies.
Bleakney (1958) suggested that the Recent population of E. blandingii
in Nova Scotia survived glaciation in an "Atlantic Coastal Plain refuge" and then
dispersed northward up the coast into Canada and Maine. Preston and McCoy
(1971), suggested that "colonization of the Atlantic Coastal Plain from the Great
Lakes region, along a 'steppe corridor' (Schmidt 1938) through the Mohawk
Valley" was a more plausible hypothesis. Preston and McCoy (1971) also stat-ed
that the study of specimens from the eastern limits may provide an answer to
the possibility of a "minor Atlantic Coast refuge for Emydoidea during ice
advances." The presence of numerous E. blandingii at the Ardis site, as well as
fossil material from New Trout Cave in West Virginia (Holman and Grady 1987),
suggest that the Recent extreme northeastern populations may be products of a
Turtles 2 9
re-invasion of the northeast by a substantial Atlantic Coastal Plain stock that had
spread at least 900 km southward during the glacial advance of the late Pleis-tocene.
Evidence from the Ardis local fauna indicates significant shifts longitu-dinally
in the spatial distribution of E. blandingii along the Atlantic Coastal
Plain during the late Pleistocene.
ACKNOWLEDGMENTS - We wish to express our gratitude to the staff of the
Giant Cement Plant, Harleyville, South Carolina, for their generous cooperation,
and in particular to Burt Ardis, for whom the fauna is named. Many thanks go
to all the field volunteers: Vance McCollum, Linda Eberle, Craig and Alice
Healy, Derwin Hudson, Ray Ogilvie, Lee Hudson, Tom Reeves, Martha Bentley,
and Karin Knight. Robert Weems, U.S. Geological Survey, and Peter Meylan,
Eckerd College, Petersburg, Florida, provided helpful discussion and review of
this manuscript. We also wish to express our appreciation to Overton Ganong of
the South Carolina State Museum; David Webb, Gary Morgan, and David Auth,
of the Florida Museum of Natural History; Dennis Hermann, of Zoo Atlanta; and
Greg Schneider, Museum of Zoology, The University of Michigan, and Justin
Cougdon, Savannah River Ecology Labratory, Aiken, South Carolina, for allow-ing
us access to their collections. Thanks go to Mike and Debi Trinkley, the
Chicora Foundation, Columbia, South Carolina, for the use of their field equip-ment,
and to Mike Runyon of Lander College, Greenwood, South Carolina, for
donating fossil material for C14 dating. Darby Erd provided the illustrations.
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Meylan, P. A. 1995. Pleistocene amphibians and reptiles from the Leisey Shell
pit, Hillsbourough County, Florida. Bulletin of the Florida Museum of Nat-ural
History 37 (9):273-297.
Mickelson, D. M., L. Clayton, D. S. Fullerton, and H. W. Borns, Jr. 1983. The
late Wisconsin glacial record of the Laurentide ice sheet in the United States.
Pages 3 1 1-353 in Late Quaternary environments of the United States. (H. E.
Wright, Jr., editor). Volume 1, University of Minnesota Press, Minneapolis.
Milstead, W. W. 1969. Studies on the evolution of box Turtles (Genus Ter-rapene).
Bulletin of the Florida State Museum 14(1): 1-1 13.
32 Curtis C. Bentley and James L. Knight
Preston, E. R. 1979. Late Pleistocene cold-blooded vertebrate faunas from the
mid-continental United States. 1. Reptilia: Testudines, Crocodilia. Universi-ty
of Michigan, Museum of Paleontology. Papers on Paleontology 19:1-53.
Preston, R. E., and C. J. McCoy. 1971. The status of Emys twentei Taylor (Rep-tilia:
Testudinidae) based on new fossil records from Kansas and Oklahoma.
Journal of Herpetology 5(l-2):23-30.
Pritchard, P. C. H. 1989. The alligator snapping turtle: biology and conservation.
Milwaukee Public Museum, Milwaukee, Wisconsin.
Roth, J. A., and J. Laerm. 1980. A late Pleistocene vertebrate assemblage from
Edisto Island, South Carolina. Brimleyana 3:1-29.
Schmidt, K. P. 1938. Herpetological evidence for the postglacial eastward
extension of the steppe in North America. Ecology 19:396-407.
Smith, W. P. 1957. An analysis of Post-Wisconsin biogeography of the Prairie
Peninsula region based on distributional phenomena among terrestrial ver-tebrate
populations. Ecology 38:205-218.
Tushingham, M. A., and W. R. Peltier. 1993. Implications of the radiocarbon
timescale for ice-sheet chronology and sea-level change. Quaternary
Research 39:125-129.
Ward, L. W., B. W Blackwelder, G. S. Gohn, and R. Z. Poore. 1979. Strati-graphic
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Received 3 January 1996
Accepted 13 August 1996
Turtles 3 3
APPENDIX I
Taxa and Minimum Number of Individuals Present
Taxa Minimum number of individuals
Kinosternon subrubrum* 5
Sternotherus odoratus* 4
Chelydra serpentina 3
Macroclemys temminckii* 1
Clemmys guttata 19
Clemmys muhlenbergii* 3
Chrysemys picta* 25
Deirochelys reticularia* 1
Trachemys scripta 4
Pseudemys sp. 1
Terrapene Carolina major 12
Emydoidea blandingii* 16
Hesperotestudo crassiscutata 1
Apalone sp. 1
Number of turtle species = 14
Number of individuals = 96
* = first fossil report from South Carolina
Observations of Freshwater Jellyfish, Craspedacusta sowerbyi
Lankester (Trachylina: Petasidae), in a West Virginia Reservoir
Ted R. Angradi
U.S. Department ofAgriculture, Forest Service
Timber and Watershed Laboratory,
Parsons, West Virginia 26287
ABSTRACT. — A swarm of medusae of the freshwater jellyfish
Craspedacusta sowerbyi was observed in a cove of a West Virginia
reservoir in August and September, 1995. Medusae were abundant
(>1000/iTr) but extremely localized. Distribution of medusae in the
cove did not appear to be linked to water chemistry. Size of medusae
ranged from 6-21 mm in diameter and increased significantly with dis-tance
from the center of abundance, suggesting that the localized distri-bution
of medusae resulted from dispersion rather than from environ-mentally-
induced aggregation. Measurements of mean diameter of
medusae on separate dates indicated a growth rate of about 0.2 mm/d,
and a medusa life cycle of approximately 102 days.
The freshwater jellyfish Craspedacusta sowerbyi Lankester 1880 is an
exotic species first observed (as medusae) in the United States in 1908 (Kramp
1950, Pennak 1989). Native to the Yang-tse River system in China (Kramp
1950), C sowerbyi has been reported from many localities worldwide between
45° north and 45° south latitude (Acker and Muscat 1976, Pennak 1989). True
freshwater jellyfishes are few, limited to about a dozen species worldwide
(Hutchinson 1967, Pennak 1989).
Craspedacusta sowerbyi has been reported from 3 1 states and the Dis-trict
of Columbia; it has not been reported from northern New England, the
Northern Rocky Mountains, or the Northern Great Plains (DeVries 1992). In
West Virginia there are records of C. sowerbyi from Barbour, Fayette, Mercer,
Monogalia, Wayne, and Wood counties (Reese 1940, Lytle 1962, Koryak and
Stafford 1981, and D. Tarter, Marshall University, personal communication).
Craspedacusta sowerbyi has two life stages, a free-swimming medusa
(10-20 mm diameter), and a sessile hydroid polyp (1 mm long, Acker and Mus-cat
1976). Lytle (1959) reviewed the developmental biology of the species.
Medusae of C. sowerbyi appear sporadically in lentic and even less frequently in
lotic ecosystems in the United States (Acker and Muscat 1976, Beckett and
Turanchik 1980, DeVries 1992). Usually a swarm of medusae appears in sum-
34
Jellyfish 35
mer where it has never before been observed or where it has not been observed
for many years (Slobodkin and Bossert 1991). Lentic systems in which the
medusae have been observed include reservoirs, natural lakes, ponds, quarries,
ornamental pools, and aquaria.
Specific environmental factors associated with the formation of
medusae from polyps via asexual reproduction (budding) are poorly understood.
Factors suggested include increasing water temperature (McClary 1959)
increased alkalinity (Koryak and Clancy 1981, McCullough et al. 1981),
increasing dissolved CO2 (Acker and Muscat 1976), decreasing stream flow
(Brussock et al. 1985), changing reservoir levels (Deacon and Haskell 1967) and
increasing supply of zooplankton (Lytle 1959), on which the medusae prey.
Dispersal of C. sowerbyi among water bodies probably occurs via
polyps attached to aquatic plants or waterfowl, or in tanks used to transport fish
(Byers 1945, Bushnell and Porter 1967, Howmiller and Ludwig 1970). The
polyps can survive in moving water (Hutchinson 1967), so once the polyps enter
a river system, the medusae may eventually appear in downstream reservoirs
(e.g., Yeager 1987).
Because the medusae occur unpredictably and the polyps are micro-scopic
and easily overlooked, the complete geographic distribution and ecology
of C. sowerbyi are not well known. Field studies have been mostly descriptive
(e.g., Garman 1916, Deevy and Brooks 1943, Dexter et al. 1949, Chadwick and
Houston 1953, Bushnell and Porter 1967, Koryak and Clancy 1981, McCullough
et al. 1981, Dodds and Hall 1984). Deacon and Haskell (1967) examined diel
activity patterns of medusae at Lake Mead, Nevada. Dodson and Cooper (1983)
examined trophic relationships of the medusae in the laboratory. Acker and
Muscat (1976) and DeVries (1992) reviewed the literature on the ecology of C.
sowerbyi.
The purpose of this paper is to describe a swarm of C. sowerbyi
medusae I observed at Stonewall Jackson Lake, Lewis County, West Virginia, in
August and September 1995. My initial observations of the medusae at
Stonewall Jackson Lake suggested that the size distribution of medusae varied
with distance from the main concentration of medusae (swarm). I hypothesized
that the distribution of medusae resulted from dispersion from the apparent pop-ulation
center at the swarm, and I predicted that medusae collected away from
the main swarm location would be larger than medusae within the swarm
because more distant medusae would have had more time to grow. The null
hypothesis that medusae collected from all locations have the same size class dis-tribution
implies that the dense concentration of medusae at the swarm location
results primarily from aggregation due to water chemistry, temperature, food,
current, wind, or some other factor rather than from dispersion. I collected and
measured specimens to test this hypothesis. I also compared the mean size of
36 Ted R. Angradi
specimens collected on separate dates to calculate an approximate growth rate
for the medusae.
METHODS
I observed the medusae at Wolf Fork (80°28'W, 38°59'N), a cove
formed by a flooded tributary to Skin Creek which forms the east arm of
Stonewall Jackson Lake (Fig. 1). Wolf Fork is about 1.5-km long and 30-150-m
wide and has abundant flooded timber. The cove is well sheltered from winds
and is a no-wake boating zone. A culvert connects Wolf Fork and the stream
draining the upper watershed. Stonewall Jackson Lake is a 1,070-ha Army
Corps of Engineers reservoir filled in 1986. The main West Fork River arm of
the reservoir is a tributary of the Tygart River in the Ohio River drainage.
During my initial visit to Wolf Fork (16 August 1995) I estimated the
density (number m3
) of medusae using two methods. Where the medusae were
abundant I used a 20-L plastic bucket. From a small boat I slipped the bucket
into the water and withdrew it with minimal turbulence. I poured the bucket con-tents
through a fine sieve and transferred the medusae to a tray for enumeration.
Where the medusa were scarce, I estimated the density visually. Both methods
are biased toward the upper 0.5 m of water surface because I could not see or
sample medusa at greater depths. On the first visit to Wolf Fork I collected water
samples and recorded water temperature and dissolved oxygen at several loca-tions
in the cove. Water samples were analyzed at the U.S. Forest Service Tim-ber
and Watershed Laboratory, Parsons, West Virginia.
Fig. 1 Map of Stonewall Jackson Lake showing the swarm location at Wolf
Fork. Inset map shows location of the reservoir in West Virginia.
Jellyfish 37
On subsequent visits to Wolf Fork (16 August and 12 September 1995)
I collected medusae with an aquarium dip net from a small boat. I measured bell
diameter of live specimens under a dissecting microscope by placing a ruler
under a clear plastic petri plate containing a few medusae and a small amount of
water. I judged measurement error to be ±1.0 mm.
RESULTS
On 16 August 1995, I observed a dense swarm of medusae near the
head of Wolf Fork. Medusae decreased greatly in abundance with distance from
the swarm. Density of medusae in three bucket samples was 1.2, 1.9, and 4.8
medusa/L. This is approximately equivalent to 1,000-5,000 medusae/m3 in the
upper 0.5 m of the water column within an area of about 25 m2
. At 100-200 m
from the swarm (toward the main channel of the reservoir) there were 10-50
medusae/m3
; at 300-400 m medusae were scarce (<1 medusa/m3
). I did not
observe medusae in lower Wolf Fork, the main channel, or in other coves of Skin
Creek, although I did not make an exhaustive search. My conversations with
anglers, reservoir managers, and local fish biologists suggest that this is the first
record of C. sowerbyi at Stonewall Jackson Lake. The origin of C. sowerbyi in
the drainage is unknown.
Table 1. Selected water quality measurements for Wolf Fork, 16 August 1995.
Site distances are from the swarm toward the main channel. All values for sam-ples
collected at the water surface. DO=dissolved oxygen.
Site Temp DO PH Conductivity Alkalinity so4 Ca Medusa
(m) C (mg/L) nSlcm mg/L
(CaCO,)
(mg/L) (mg/L) abundance
(#/m
13 D
Jfc=-;
Cane Rfver jj.
\ North Toerfiiver
vt\
'w J
/'jm
-^
—
^-r ) L
A South Toe River
1N
) J
1 1 1
NORTH CAROLINA \f 16 32
KILOMETERS
METHODS AND MATERIALS
We sampled 99 sites in 105 collections: 32 sites in the lower reach of
the French Broad River and some of its tributaries from May 1994 to May 1995,
and at 2 sites in the middle reach of the river in September 1993 (Fig. 4); 39 sites
in the Nolichucky River system between July 1991 and November 1994 (Fig. 4);
and 15 sites in the Dan River and in 2 of its tributaries 25 times from July 1992
to May 1995 (Fig. 3). We sampled most sites only once, although four sites each
in the French Broad and Nolichucky rivers were each sampled twice. We also
sampled 1 1 sites in the Virginia portion of the upper Dan River from November
1993 to October 1994 (Fig. 3). All site locations and dates are available from the
senior author. In addition, we include data from 146 collections made by others,
other personal communications, and from the literature.
We sampled primarily with a backpack electroshocker and seine, using
the technique described by Jenkins and Burkhead (1975). Each site was elec-trofished
for 45- 190 minutes, i.e., until we believed that sampling had been com-
Distribution of Fishes 4 9
prehensive. However, in the French Broad and in the Nolichucky rivers, we sam-pled
12 sites in 1991 and 2 in 1994 only by seine (3.05 m x 1.2 m, 0.64 cm mesh),
and we sampled 2 sites in the former river in 1994 only by a 25 m, 14 cm
stretched mesh monofilament gill net and a 50 m, 5.1 cm stretched mesh
monofilament gill net. The nets were deployed overnight and fished on consec-utive
days for a total of six net days. In the Dan River in 1992 and in 1994, we
sampled two sites in each year only by seine (size as above). At five sites in the
North and South Toe rivers, in addition to sampling with electroshocker and
seine, we also surveyed fishes underwater by snorkeling. In all sampling efforts,
the known preferred habitat for each species was sampled most intensively.
In addition to the fishes taken, data on stream depth, width, and sub-strate
type; current; air and water temperatures; pH; and dissolved oxygen con-centration
were often recorded at a site, and we include these data when avail-able.
Fishes were preserved in 10% formalin upon capture for subsequent exam-ination.
Fish measurements when available are given; TL is total length and SL
is standard length. We deposited preserved specimens in the North Carolina
State Museum of Natural Sciences in Raleigh. Scientific and common names of
fishes used herein follow Mayden et al. (1992), except for Cyprinella monacha
which follows Jenkins and Burkhead (1994).
We include figures that show all of our known fish capture localities in
North Carolina. In our species accounts we occasionally include records of fish-es
taken in portions of adjacent states in an effort to make the accounts more
accurate and complete.
Positive results are gratifying, but, as usual, negative results are not nec-essarily
conclusive. When fish populations in rivers and large creeks decrease
strongly, it becomes virtually impossible to differentiate between occurrence at a
low level and extirpation (Etnier 1994).
RESULTS AND DISCUSSION
TENNESSEE RIVER DRAINAGE
ENDANGERED SPECIES
Paddlefish, Polyodon spathula (Walbaum)
The paddlefish once occurred throughout the Mississippi River and its
larger tributaries, but its distribution has decreased coincidental with river chan-nelization,
damming, and overfishing (Burr 1980). Cope (1870) maintained that
it migrated up the French Broad River as far as Asheville in Buncombe County,
North Carolina. Fishermen in this state reported that it had been caught in the
lower reaches of the French Broad River as recently as 1983, but none of these
reports has been substantiated by specimens (E. Menhinick, personal communi-cation,
1994). We sampled with large-mesh gill nets in the French Broad River
downriver of Hot Springs and in the river at the mouth of Big Laurel Creek, both
Madison County, North Carolina, on 14 and 15 May and 15 and 16 August, 1994,
50 Fred C. Rohde, Mary L. Moser and Rudolf G. Arndt
respectively. Swift currents limited our efficiency. We caught no paddlefish.
Local fishermen in Madison County whom we questioned in 1994 told us that
they had never seen a paddlefish from North Carolina nor heard of one caught
there. We presume that the paddlefish has been extirpated from North Carolina.
Fig. 5. Distribution of the stonecat, Noturus flavus, (circle) and the blotchside
logperch, Percina burtoni, (star) in the French Broad and Nolichucky river sys-tems,
North Carolina. An open circle overlaps two historical sites where the
stonecat was not taken in this survey. Specific historical sites for the blotchside
logperch in Cane Creek and the Swannanoa River are not known and are plotted
as circles with a question mark.
Stonecat, Noturus flavus Rafinesque
The stonecat is distributed through portions of the Mississippi River
basin, the Great Lakes, the Ohio River basin, and the St. Lawrence, Mohawk, and
Hudson River systems (Rohde 1980). In North Carolina it is documented from
only three sites in the Cane River, where it was collected on 1 8 June (one speci-men)
and 26 June (six), 1984, and 15 September 1985 (two) (Menhinick 1986).
Despite our efforts to collect it at all three sites, we took two adults (pho-tographed
and released) only at the downstream-most site on 4 September 1993
(Fig. 5).
Distribution of Fishes 5 -j
We record it here for the first time from the Ivy River, a tributary to the
French Broad River, from upstream of Marshall in Madison County, North Car-olina,
where we took three adults (90-99 mm SL) on 14 August 1994 (Fig. 5).
One adult (86 mm TL) and one specimen (released, not measured) were collect-ed
in the Little Tennessee River at Needmore in Swain County, North Carolina,
by Tennessee Valley Authority (TVA) personnel on 20 June 1994 (E. Scott, per-sonal
communication, 1994) (Fig. 1). This site had been sampled five times by
TVA biologists during the period 1988-1993 and once by us in 1993. The dis-covery
of the species there in 1994 was unexpected. Preferred habitat was grav-el
riffles. Current at the Ivy River site was 0.31 m/sec, water temperature 20.6
C, pH 8.0, and dissolved oxygen concentration 8.0 ppm. Its status of endangered
in North Carolina is warranted.
Blotchside logperch, Percina burtoni Fowler
The blotchside logperch occurs in disjunct populations in the Ten-nessee
River drainage from westcentral Tennessee to southwestern Virginia
(Page and Burr 1991); where it occurs it is localized and rare (Etnier 1994). In
North Carolina it was taken at one site in Cane Creek (Henderson County) in
1902, at one site in the Swannanoa River (Buncombe County) in 1934, and at
two sites in the South Toe River (Yancey County) in 1975 and 1977 (Menhinick
1986) (Fig. 5). We collected two adults (120 mm SL, one released) at a new
locality in the South Toe River near its confluence with the North Toe River in
September 1993. Since we noted that this fish can readily avoid electroshockers
and seines, we observed 2 to 6 adults in each of 4 visits by snorkeling at the two
upstream historic sites in the South Toe River in July and September 1993 and
August 1995 (Fig. 5). Preferred habitat was in pools below riffles. Menhinick
(1986) presumed P. burtoni to have been extirpated from the Swannanoa River
and from Cane Creek since he did not obtain it there in 14 collections nor did the
North Carolina Division of Environmental Management in 5 collecions (V
Schneider, personal communication, 1994). We did not collect it in either stream
in two collections made there in 1993, and we concur with Menhinick (1986). Its
continued presence in North Carolina is tenuous.
Dusky darter, Percina sciera (Swain)
The dusky darter occurs from the Wabash River drainage in Indiana
south and west to the Guadalupe River drainage in Texas and east to the Tombig-bee-
Black Warrior river system in Alabama (Page 1980). In North Carolina it is
known only from Spring Creek, Madison County, where U.S. Forest Service per-sonnel
collected one specimen in 1966 and one in 1969 (Auburn University Col-lection
3442), although the specific sites are now not known (M. Seehorn, per-sonal
communication, 1994). We made 13 collections at 7 sites with suitable
habitat in Spring Creek over a distance of 27.3 rkm in 1994 and 1995. We did
not collect it. Apparently never widespread or common in North Carolina, we
52 Fred C. Rohde, Mary L. Moser and Rudolf G. Arndt
consider the dusky darter to have been extirpated from this state by unknown
causes. It appears to be a relatively tolerant species in other portions of its range.
THREATENED SPECIES
American brook lamprey, Lampetra appendix (DeKay)
The American brook lamprey is widely distributed in the St. Lawrence
and Mississippi river basins from New York to northern Arkansas, and on
Atlantic Slope drainages from southern Quebec south to the Roanoke River
drainage in Virginia (Page and Burr 1991). In North Carolina it was known from
only one site in the downstream reach of Spring Creek, Madison County, at a
point where a railroad trestle crosses this creek, where 26 individuals were taken
in 1980 and 1 in 1983 (Menhinick 1986). We took one adult some 200 m down-stream
of the above site (and about 50 m upstream of the confluence of Spring
Creek with the French Broad River, Madison County) on 22 April 1995 (Fig. 6),
and two adults on the same day, also in Spring Creek at a point about 0.9 rkm
above this confluence. They were males and measured 144, 147, and 157 mm
TL. All were taken in gravel riffles where the current was 0.45 m/sec. We col-lected
one ammocoetes of 152 mm TL in a sandy-bottomed pool about 50 m
downstream of the last-mentioned Spring Creek site on 14 August 1994. The pH
here was 6.9, and the dissolved oxygen concentration was 8.4 ppm. This species
appears to be restricted to this creek in North Carolina. Its status of threatened
in North Carolina appears to be conservative.
Fig. 6. Distribution of the American brook lamprey, Lampetra appendix, (circle)
and the striped shiner, Luxilus chrysocephalus, (star) in the French Broad and
Nolichucky river systems, North Carolina. Some symbols overlap sites.
NORTH CAROLINA
Distribution of Fishes 5 3
Spotfin chub, Cyprinella monacha (Cope)
The spotfin chub is endemic to the Tennessee River drainage in disjunct
populations from southwestern Virginia to northwestern Alabama and in the Buf-falo
River in central Tennessee (Jenkins and Burkhead 1984, Etnier and Starnes
1994). It has apparently been extirpated from Alabama and Georgia (Etnier and
Starnes 1994). In North Carolina it is restricted to a 16.8 rkm section of the Lit-tle
Tennessee River in Macon and Swain counties (Alderman 1987). We col-lected
and released 27 spotfin chub in this river at Needmore in Swain County
(Fig. 1) on 25 September 1993. Three previous North Carolina records, two
from the French Broad River system (1888) and one from the Tuckaseegee River
(1940), apparently represent now extirpated populations (Menhinick 1986). We
did not take it in 36 collections in the French Broad River system, and we con-cur
with Menhinick (1986) that it has been extirpated from this drainage. Its sta-tus
of threatened in North Carolina appears to be conservative.
Striped shiner, Luxilus chrysocephalus Rafinesque
The striped shiner is common in the southern Great Lakes basin from
western New York and southeastern Wisconsin south through much of the Mis-sissippi
River basin almost to the Gulf of Mexico (Page and Burr 1991). A dis-junct
population was discovered in the Cane River, Yancey County, North Car-olina
in 1980 by E. Menhinick, and soon thereafter it was known from five sites
in the Cane River system (Menhinick 1986). We found it in four sites in 14.5
rkm of the middle reach of the Cane River and in two tributaries, Bald and Indi-an
creeks (Fig. 6), in 1994. Numbers taken per our collections ranged from 2-
23. Specimens ranged from 29-107 mm TL. Tennessee Valley Authority biolo-gists
took 61 individuals in one collection in the Cane River in 1992 (E. Scott,
personal communication, 1994) (Fig. 6). Preferred habitat was pools and runs.
Current ranged from 0.39-0.57 m/sec, pH 7.0-7.5, and dissolved oxygen concen-tration
6.8-8.4 ppm. Its status of threatened in North Carolina is warranted.
Banded sculpin, Cottus carolinae (Gill)
The banded sculpin inhabits mountainous areas of the Mississippi River
basin from West Virginia west to Kansas and from the Ozark Mountains south-east
to southern Alabama (Page and Burr 1991). In North Carolina it was report-ed
only from Big Laurel and Spring creeks, Madison County (Robins 1954).
Menhinick (1986) later reported it as restricted to the main stream of the French
Broad River in North Carolina near the Tennessee line and absent from the two
creeks. E. Menhinick (personal communication, 1994) took two adults and eight
juveniles with rotenone in the downstream-most 100 m of Shut-in Creek, Madi-son
County, North Carolina in July 1994 (Fig. 7). We did not collect it at two
upstream-sites in this creek in 1994. However, we did take 58 specimens in two
collections made on 14 May and 19 July 1994 throughout the lower 300 m of
54 Fred C. Rohde, Mary L. Moser and Rudolf G. Arndt
Paint Creek, Greene County, Tennessee. This creek enters the French Broad
River some 120 m downstream of the North Carolina/Tennessee line (Fig. 7). Its
status as threatened appears to be conservative.
Fig. 7. Distribution of the banded sculpin, Cottus carolinae, (circle) and the
freshwater drum, Aplodinotus grunniens, (star) in the French Broad River sys-tem,
North Carolina. A circle with a question mark indicates an undefined his-torical
site of the banded sculpin. Some symbols overlap sites.
Logperch, Percina caprodes (Rafinesque)
The logperch occurs from central Canada and the upper Mississippi
River and adjacent drainages south to the Gulf of Mexico, and on Atlantic Slope
drainages from the Hudson River south to portions of the Chesapeake Bay
drainage (Rohde et al. 1994). In North Carolina it is known from four sites in
the French Broad River between Redmon Dam and the Tennessee state line
(Harned 1979) (Fig. 8), four specimens were collected below Redmon Dam in
1986 and 1987 (Birchfield et al. 1987) (Fig. 8), and from one site in the New
River, Allegheny County (Menhinick 1986). We collected eight adults (88-132
mm SL) at four sites in the downstream reaches of the French Broad River and
at two sites in the downstream portion of Spring Creek, Madison County on 13
May; 19, 22 July; and 5 November 1994 (Fig. 8). One specimen was taken in
Distribution of Fishes 55
Fig. 8. Distribution of the logperch, Percina caprodes, (circle) and the olive
darter, Percina squamata, (star) in the French Broad and Nolichucky river sys-tems,
North Carolina. Some symbols overlap sites.
cd^ ^W^^ TENNESSEE / /'
N—I
WQJj/^hi ir*ir-iy Diuar
\^£.s-~
S -\ f**
French Broad River -*~^~,~}\
Cane River j
1 North Toeffiiver
vV*w \
r"J\
-*/—^-r ) F l/_
) r" \
/* South Toe River
1N
1 J
NORTH CAROLINA \f 1 6 32
KILOMETERS
Spring Creek 7.1 rkm upstream from its mouth in July 1994 (S. Bryan, personal
communication, 1994) (Fig. 8). Preferred habitat in the river was runs with large
boulders. Current at our Spring Creek site was 0.58 m/sec, pH 7.1-7.6, and dis-solved
oxygen concentration 7.1-10.5 ppm. Its status of threatened is warranted.
Freshwater drum, Aplodinotus grunniens Rafinesque
The freshwater drum occurs throughout the Mississippi River basin
from southern Canada and the Great Lakes to western Texas and western Flori-da
(Rohde et al. 1994). Prior to this survey, it was known in North Carolina from
six sites in the lower reaches of the French Broad River downstream of Redmon
Dam, Madison County (Harned 1979) (Fig. 7). We collected one large specimen
(305 mm TL) in a pool in Spring Creek, at a point 1 rkm upstream of its conflu-ence
with the French Broad River, on 22 July 1994, and E. Menhinick (personal
communication, 1994) took one specimen in the same month in Spring Creek at
this confluence (Fig. 7). Its status of threatened is warranted due to the lack of
juveniles in collections.
5 6 Fred C. Rohde, Mary L. Moser and Rudolf G. Arndt
SPECIES OF SPECIAL CONCERN
Lake sturgeon, Acipenserfulvescens Rafinesque
The lake sturgeon is usually found over shoals in lakes and large rivers
in central Canada and Hudson Bay and St. Lawrence River drainages, and in
much of the Mississippi River drainage south to northeastern Louisiana (Page
and Burr 1991). Eight specimens, presumably of this species, were taken from
the French Broad River near Hot Springs in Madison County, North Carolina in
1945 (Brimley 1946). An occasional lake sturgeon is still reported from Douglas
Reservoir in Jefferson County, Tennessee, but these are unsubstantiated records
(Etnier and Starnes 1994). We set 2 large-mesh gill nets of 25 and 50 m total
length in the French Broad River downstream of Hot Springs in mid-May 1995
and in the river at the mouth of Big Laurel Creek in mid-August but failed to col-lect
sturgeon. Swift current limited sampling location possibilities at the former
site and reduced gear efficiency. Local North Carolina state fishery biologists
have no reported sightings (J. Borawa, personal communication, 1994). Men-hinick
(1986) considers the lake sturgeon to have been extirpated from North
Carolina, and we concur.
Mooneye, Hiodon tergisus Lesueur
The mooneye is found in central and southern Canada and in much of
the Mississippi River basin from the Great Lakes south to the Gulf of Mexico
(Page and Burr 1991). It historically occurred in the upper reaches of the French
Broad River near Bowman's Bluff, Henderson County, North Carolina in 1902
(Smith 1907), but it is now known only from Redmon Dam to the Tennessee
state line (Menhinick 1986) based on several mooneye obtained from fishermen
in the French Broad River just above the confluence with Big Laurel Creek by
Harned (1979). We did not take it in this river in our electroshocker or gill net
collections. Its status of special concern in North Carolina appears to be conser-vative.
River carpsucker, Carpiodes carpio (Rafinesque)
The river carpsucker occurs throughout the Mississippi River basin
from Montana to Pennsylvania and south to the Gulf of Mexico (Lee and Plata-nia
1980). There is one North Carolina 1947 record from the French Broad River
near Hot Springs in Madison County (Menhinick 1986). It was also captured in
the same river in Tennessee 41 rkm downstream of the North Carolina state line
in 1979 (Harned 1979), but we failed to collect it in this study. Its status of spe-cial
concern appears to be conservative.
Mountain madtom, Noturus eleutherus Jordan
The mountain madtom occurs in disjunct populations from northwest-ern
Pennsylvania south and west through the Ohio River basin to the Red and
Distribution of Fishes 57
Ouachita river drainages in Oklahoma and Arkansas (Page and Burr 1991). The
only verified North Carolina specimens are from Spring Creek, Madison Coun-ty
and were collected in 1889 (Taylor 1969). It was also collected at two sites in
the French Broad River just upstream of Douglas Reservoir, Cocke County, Ten-nessee
(32 km downriver of North Carolina) during a 1979 TVA survey (Harned
1979). We did not collect it at any of the 34 sites we surveyed in the lower reach-es
of the French Broad River system. We concur with Menhinick (1986) that it
has been extirpated from North Carolina.
Snubnose darter, Etheostoma simoterum (Cope)
The snubnose darter is abundant in the Tennessee River drainage from
southwestern Virginia to northern Alabama (Rohde et al. 1994). The only puta-tive
extant specimen from North Carolina was reported by Cope (1870) and is
now in the United States National Museum, but it is unclear from Cope's records
whether its provenance is North Carolina or Tennessee (Menhinick 1986). Men-hinick
(1986) reported two unverified records from Laurel and Spring creeks,
Madison County, North Carolina. We collected no snubnose darter, nor did Men-hinick
(1986). M. Hopey, who made 13 collections for a general survey of the
streams in this area for the Western North Carolina Alliance in 1992, did not col-lect
it (M. Kelly, personal communication, 1994). We consider the past or pre-sent
occurrence of this darter in North Carolina to be highly doubtful.
Wounded darter, Etheostoma vulneratum (Cope)
The wounded darter is restricted to the upper Tennessee River drainage
from Virginia to Georgia (Rohde et al. 1994). It is abundant in the Little Ten-nessee
River in North Carolina (F. Rohde, personal observations). Although the
type locality is Spring Creek, Madison County, North Carolina (Cope 1870),
none has been reported from the French Broad River system in North Carolina
since then, including our survey. Harned (1979) collected one specimen in the
French Broad River in Tennessee at a point approximately 23 km downstream of
the North Carolina state line. We conclude that it has been extirpated from the
French Broad River system in North Carolina. Its status of special concern in
North Carolina appears conservative.
Olive darter, Percina squamata (Gilbert and Swain)
The olive darter is confined to the Rockcastle and Big South Fork rivers
in the Cumberland River drainage in Kentucky and Tennessee and to the upper
Tennessee River drainage (Rohde et al. 1994). There are five records from the
lower reaches of the French Broad River system (three in the main river and two
in Spring Creek) in North Carolina, and four records from the Nolichucky River
system (three in Cane River and two in North Toe River); it also occurs in the
Little Tennessee and upper Hiwassee rivers in the state (Menhinick 1991). We
58 Fred C. Rohde, Mary L. Moser and Rudolf G. Arndt
did not collect it in the French Broad River system, but we collected one juve-nile
(38 mm SL) in the Cane River on 19 July 1993, and three adults (104 mm
SL, two released) in the South Toe River on 5 September 1993 (Fig. 8). Pre-ferred
habitat is around large boulders in fast riffles. Its status of special concern
in North Carolina appears to be conservative.
NEW STATE RECORD
Ohio lamprey Ichthyomyzon bdellium (Jordan)
The Ohio lamprey occurs in disjunct populations in the Ohio River
basin, where it is uncommon (Rohde and Lanteigne-Courchene 1980). Because
of its presence in nearby Tennessee, Menhinick et al. (1974) listed its occurrence
in North Carolina as probable. However, there were no records from North Car-olina
until we took one male and three females from the mouth of Spring Creek,
Madison County, on 14 May 1994 (Fig. 9). Each was adult (220-260 mm TL),
had a well-developed digestive tract, and each female was gravid. We took
another three males (243-246 mm TL), one female (244 mm TL), and two juve-niles
(not ammocoetes, nor mature adults) (152, 153 mm TL) here on 22 April
1995, as well as two adult males (239, 248 mm TL) 1 rkm further upstream in
this creek on the same day. We took three females (235-240 mm TL) in nearby
Paint Creek, Greene County, Tennessee on 14 May 1995. All our specimens
were taken over rocky riffles with a current from 0.45-0.78 m/sec.
Fig. 9. Distribution of the Ohio lamprey, Ichthyomyzon bdellium, in the French
Broad River system, North Carolina.
TENNESSEE
Nolichucky River ^ -.
Distribution of Fishes 59
DAN RIVER SYSTEM
ENDANGERED
Cutlips minnow, Exoglossum maxillingua (Lesueur)
The cutlips minnow occurs on the Atlantic slope from the St. Lawrence
River and eastern Lake Ontario drainages south to the upper Dan River in North
Carolina (Gilbert and Lee 1980). Menhinick (1986) reported it from one site on
the Dan River in Stokes County, North Carolina, within 1.6 rkm downstream of
the Virginia state line. We found it in the North Carolina portion of this river at
four sites from the Virginia line downstream to NC Route 704 and at six Virginia
sites upstream to the Pinnacles Power Plant, over a total distance of 43 rkm (Fig.
10). Numbers (39) taken in our collections ranged from 1-6 (mean 3.9), and their
length ranged from 69-133 mm SL; most specimens were adults. This species
preferred fast-flowing runs or pools, near large rocks or boulders over sand and
gravel. Current where it was collected was 0.54-0.75 m/sec; water temperature
7.7 C (November)-22 C (July); pH 6.8-7.6; and dissolved oxygen concentration
10.6-11.8 ppm (both November). The species appears to be secure in its limited
distribution in North Carolina.
Fig. 10. Distribution of the cutlips minnow, Exoglossum maxillingua, in the Dan
River system, North Carolina and Virginia.
60 Fred C. Rohde, Mary L. Moser and Rudolf G. Arndt
Rustyside sucker, Thoburnia hamiltoni Raney and Lachner
The rustyside sucker is endemic to the upper Dan River system in North
Carolina and Virginia (Jenkins and Burkhead 1994). In North Carolina it is
known only from the 1 .4 rkm downstream-most portion of the Little Dan River
in Stokes County. Here Menhinick (1986) collected four specimens at a point
some 400 m downriver of the Virginia line in 1985 (Fig. 11). We made three col-lections
in the Little Dan River, from its confluence with the Dan River upstream
to the North Carolina/Virginia line, and took one adult (144 mm SL) in a run with
gravel and rubble substrate on 21 December 1992. In Virginia we took three
adults (1 18-142 mm SL) in the Dan River at one site located 365 m, and at anoth-er
site 914 m, downriver of the Pinnacles Power Plant on 28 November